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of the ^ 
Soviet Union 

VG. Heptner 
A. A. Sludskii 

The present book, part of a three- 
volume monograph, is a continuation 
of Volume II, Part 1 , which was 
devoted to sea cows and carnivores. It 
contains species descriptions of 
terrestrial carnivores and detailed 
information on their external morpho- 
logy, skull, body measurements and 
other data, affinities with other 
species, geographic distribution in the 
historic past and today, geographic 
variation, practical significance, and 
biology. Descriptions are presented 
for orders, families, and genera, and 
keys given for their identification. 

The book is richly illustrated with 
photographs, sketches, and colored 
illustrations by the famous wildhfe 
painter, A.N. Komarov, and the 
zoologist-artist, N.N. Kondakov. 

The results of original scientific 
studies are pubUshed here for the first 
time, providing readers a vast wealth 
of material heretofore unknown. This 
work is intended for teachers and 
students of faculties of biology and 
geography in universities, as well as 
pedagogic, agricultural, and forest 
institutes, similar organizations, 
workers engaged in game, fur, forest, 
and fish trades, those interested in the 
conservation of nature, and all persons 
interested in zoology and nature 

Mammals of the Soviet Union 

Volume II 
Part 2 


In Three Volumes 

Edited by 
V.G. Heptner and N.P. Naumov 

Vysshaya Shkola Publishers 
Moscow, 1972 

Mammals of the 
Soviet Union 

Volume II, Part 2 

(Hyaenas and Cats) 

V.G. Heptner and A.A. Sludskii 

A.N. Komarov and N.N. Kondakov 

Scientific Editor 
Robert S. Hoffmann 

Smithsonian Institution Libraries 


The National Science Foundation 

Washington, D.C. 


ТТ 76-52041 

Mlekopitayushchie Sovetskogo Soyuza 

In Three Volumes 

V.G. Heptner and N.P. Naumov, editors 

Vysshaya Shkola Publishers 

Moscow, 1972 

Translator: P.M. Rao 

General Editor: Dr. V.S. Kothekar 

© 1992 Amerind Publishing Co. Pvt. Ltd., New Delhi 

Library of Congress Cataloging-in-Publication Data 

(Revised for volume 2, pt. 2) 

Geptner, V.G. (Vladimir Georgievich), 1901-1975. 
Mammals of the Soviet Union. 

Translation of: Mlekopitaiushchie Sovetskogo Soiuza. 

Bibliography: v. 2, pt. 2, p. 

Vol. 2, pt. 2 has index. 

Supt. of Docs, no.: SI 1.2: Ar 7/2 

Contents: v. 1. Artiodactyla and Perissodactyla — 
V. 2. — pt. 2. Camivora (Hyaenas and cats). 

1. Mammals — Soviet Union. I. Nasimovich, A.A. 
II. Bannikov, Andrei Grigor'evich. III. Hoffmann, Robert S. 
IV. Title. 
QL728.S65G4713 1988 599.0947 85-600144 

Translated and published for the Smithsonian Institution Libraries, 

pursuant to an agreement with the National Science Foundation, Washington, D.C. 

by Amerind Publishing Co. Pvt. Ltd., 66 Janpath, New Delhi 110 001 

Printed in India at Pauls Press, New Delhi 


The present book, part of a three-volume monograph, is a continuation of 
Volume II, Part 1, which was devoted to sea cows and carnivores. It contains 
species descriptions of terrestrial carnivores and detailed information on 
their external morphology, skull, body measurements and other data, 
affinities with other species, geographic distribution in the historic past and 
today, geographic variation, practical significance, and biology. Descriptions 
are presented for orders, families, and genera, and keys given for their 

The book is richly illustrated with photographs, sketches, and colored 
illustrations by the famous wildlife painter, A.N. Komarov, and the 
zoologist-artist, N.N. Kondakov. 

The results of original scientific studies are published here for the first 
time, providing readers a vast wealth of material heretofore unknown. This 
work is intended for teachers and students of faculties of biology and 
geography in universities, as well as pedagogic, agricultural, and forest 
institutes, similar organizations, workers engaged in game, fur, forest, and 
fish trades, those interested in the conservation of nature, and all persons 
interested in zoology and nature study. 

Foreword to the English Edition 

The Smithsonian Institution Libraries, in cooperation with the Natiotial 
Science Foundation, has sponsored the translation into English of this and 
Mildreds of other scientific and scholarly studies since 1960. The program, 
funded with Special Foreign Currency under the provisions of Public Law 
480, represents an investment in the dissemination ofhiowledge to which 
the Smithsonian Institution is dedicated. 

The present volume of the series Mammals of the Soviet Union, Carnivores 
(Hyaenas and Cats), is the second volume to be published in English. 
Previously Volume I, Artiodactyla and Perissodactlya, was published in 
1988. A third English translation, Volume II, Part 3, Pinnipeds and Tooth- 
whales, is in preparation and should be published in the next year or two. 
This will leave only one part of the original Russian language series 
untranslated; namely. Volume II, Part 1, Sea cows and Carnivores (Dogs, 
Bears, and Mustelids). The fact that this latter volume was not published 
in English translation in the proper sequence is due to the fact that the 
translation was performed under a different contract from that supervised 
by P.M. Rao of Amerind Publishing Company, New Delhi, India. However, 
it is the intention of the Smithsonian Libraries Translation Program to publish 
this final volume as soon as possible. 

As indicated in the foreword to the English edition of Volume I, the 
senior author and editor of the series, Dr. Vladimir Georgievich Heptner, 
died in 1975, and did not complete the projected series. However, work 
continues, and volumes on baleen whales and lagomorphs are in progress. 
It is to be hoped that English translations of these volumes, when they appear, 
will follow more promptly than has been true of the first volumes of the 

English readers interested in Dr. Heptner's contributions as a 
mammalogist should refer to the foreword to the English edition of Volume 
I. Conventions used in rendering geographic names, first stated there, are 
reprinted here for the convenience of the reader. Geographic names are 
generally transliterated directly, but a few exceptions were permitted (e.g. 
Moscow instead of Moskva, translation rather than transliteration of certain 
modifiers of place names, such as Northern, rather than Severnaya, Dvina). 
Soviet administrative units are numerous, and the following equivalents 
were employed in translation: Krai, territory; oblast', district; raion, region; 


guherniya (archaic), province. Also, in the original Russian text, rivers, 
mountain ranges, and cities are often not explicitly identified, the Soviet 
reader being presumed sufficiently familiar with the geography of the country 
to be able to understand from the context of the sentence what sort of place 
is referred to. Complicating the matter is the lack of articles as parts of 
speech in Russian. To assist the English reader, I have adopted the following 
conventions: if a river is referred to, an article precedes it; if a mountain 
range is referred to, it is translated as a plural; if a city is referred to, it 
is singular, and lacks the article. Examples are: the Ural (river); the Urals 
(mountains); Ural'sk (city). Geographic place names are also inflected in 
Russian, and these have been simplified by omitting transliteration of the 
inflected ending. For example, the Russian phrase v Yaroslavskoi i 
Kostromskoi oblastyakh is translated "in the Yaroslavl and Kostroma 
districts." In cases where the nominative form of the place name has an 
-sk ending this is, however, transliterated (e.g., Omsk); when a Russian 
"soft sign" is employed in a place name, this is transliterated as an 
apostrophe (e.g., Khar'kov). Because of the large number of place names 
in this volume, it was not possible to verify all of them, and some 
inconsistencies are likely to occur. 1 would appreciate it if readers would 
bring any errors they may notice to my attention. 

One further point of confusion not apparent to me when Volume I was 
translated also requires clarification; that is the English transliteration of 
the senior author's sumame. This begins with the fourth letter of the Cyrillic 
alphabet, which usually has a "G" sound in Russian. However, the sumame 
was originally German (as indeed was mine) and in the original German 
began with the letter "H" of the Latin alphabet. Since Cyrillic has no 
equivalent of "H" this is usually transliterated into "G" in Russian. 
However, I know from conversations with him that Dr. Heptner preferred 
to use the original Germanic form of his sumame rather than the transliterated 
version, which is rendered as Geptner. The rules of transliteration employed 
by the Library of Congress do not permit such flexibility, and the attentive 
reader may notice that Library of Congress cataloguing employs the latter. 

June 1989 Robert S. Hoffmann 

Assistant Secretary for Research 
Smithsonian Institution 
Washington, D.C. 


5 The present book is a continuation of the second volume of a three-volume 
monograph on mammals of the Soviet Union and is devoted to a description 
of terrestrial carnivores of the superfamily Feloidea, i.e., the hyaena 
(Hyaenidae) and cat (Felidae) families, thus concluding a description of 
the order Camivora. In accordance with the scheme laid out in previous 
volumes, this work also includes a description of the lion, which lived in 
our country in the historic past. 

Considerable information has accumulated in the last decade on cats, 
partly due to current interest in this group. Populations and distribution 
of these species have rapidly shrunk and some species are on the verge 
of extinction. This has evoked particular attention in cats not only from 
a general biological point of view, but also in the context of conserving 
natural resources. On the whole, information on cats is so extensive that 
it necessitated their inclusion in a separate book, using a slightly different 
style of presentation. 

Part 3 concerns pinnipeds and cetaceans. Since it is preferable to give 
a description of Soviet marine animals in a single volume, the taxonomic 
sequence of the description of orders (pinnipeds after carnivores) had to 
be slightly modified; later, however, the general sequence in the more 
specialized orders has been maintained (see "Systematics of the Class 
Mammalia"). Within the orders, as in earlier volumes, descriptions of taxa 
are presented in the order of increasing specialization. 

All the characters of the groups and species have been given according 
to the scheme adopted in preceding volumes. A departure has been made 
only in the case of lions, and the silvestris-libyca, which are quite complex 
taxonomic examples of semispecies or superspecies. Paleontological data 
have been briefly summarized, mainly from the works of Simpson (1945) 
and Thenius and Hofer (1960), and in some cases other more specialized 

As in earlier volumes, the range descriptions are generalized; only the 
boundaries are described, based on an analysis of all available data. Only 
major reference points are indicated for the periphery. As far as possible, 
historical changes in the range have been traced and, on their basis, 
"reconstructed" ranges restricted to the historic period established. In spite 
of all the suppositions involved in such an approach, it is the only way 
to establish a fairly natural range. A knowledge of the latter is an essential 

prerequisite not only for resolving theoretical aspects of zoogeography, but 
also for field work for different fauna. ^ 

The reconstructed range outside the Soviet Union has been treated in 
very general terms and based on a large number of special studies, but mainly 
the critically verified accounts of Miller (1912), Sowerby (1923), Anthony 
(1928), Aharoni (1930), Nezabitovskii (1934), Lukashkin and Zhemakov 
(1934), Shortridge (1934), Phillips (1935), Heim de Balzac (1936), Allen 
(1938-1940, 1939, and 1942), Pocock (1934 and 1941), Cabrera and Yepes 
(1940), Chasen (1940), Hamilton (1943), Harper (1945), Simpson (1945), 
Anderson (1946), Carter, Hill and Tate (1946), Prater (1947), Tate (1947), 
Troughton (1948), Anar* (1949), Kalinesku (1951), Roberts (1951), EUerman 
and Morrison-Scott (1951 and 1966), EUerman, Morrison-Scott and Hayman 
(1953), Bannikov (1954), Laurie and Hill (1954), Miller and Kellog (1955), 
Kosswig (1955), Siivonen (1956 and 1967), Haltenorth and Trenze (1956), 
van den Brink (1958), Frechkop (1958), Cabrera (1958 and 1960), Shaw 
(1955 and 1958), Markov (1959), Misonne (1959), Murie (1959), Hall and 
Kelson (1959), Hatt (1959), Dulic and Tortric (1960), Vasiliu (1961), Burt 
and Grossenheider (1962), Burton (1962), Cockrum (1962), Atanasov and 
Peschev (1963), and several others. Special references have not been made 
to these works in the text. The North American distribution given in the 
maps, in almost all cases, is based on the data of Hall and Kelson (1959). 

The morphological characters of the species described are mostly based 
on the latest data gathered in the last decade in the Zoological Museum 
of Moscow University. These have made possible a fresh revision of the 
geographic variation of all species. For parts of the range outside the Soviet 
Union the list of geographic forms has usually been given without comment, 
or only with some general remarks. 

All synonyms, as in previous volumes, have been worked out anew 
and given in strictly limited numbers on the basis of the following principles: 
only true synonyms have been mentioned; from among generic synonyms 
only those for which species of Russian fauna serve as type species, or 
if the generic name is of importance, or is sometimes used (or was 
previously), or could be applied to Russian species. All names, prevailing 
or otherwise, have been given among synonyms of species in the following 
instances: 1) if the name is from a nominate species from Soviet territory; 
2) if the name is based on a nominate species from other territories, but 
is of importance and is used or was previously for one or another Russian 
form; 3) if the name is based on animals from other territories and not 

'For more details on the reconstructed range, see V.G. Heptner, Dinamika areala 
nekotorykh kopytnykh i antropokul'turnyi faktor (Range dynamics of some ungulates and 
the anthropocultural factor). Voprosy Geografii, 1960, vol. 48. Moscow. 

*Not clear; not in Literature Cited — Sci. Ed. 


applied to Russian forms, but could be of importance for some of them; 
and 4) the name of the nominate form is invariably given irrespective of 
its type locality, whether falling within or outside Soviet boundaries. The 
actual names have not been shown separately among the synonyms of 
species, but are given in the list of subspecies inhabiting the Soviet Union 
and the synonyms pertaining to a given subspecies likewise are listed there. 
Synonyms of subspecies inhabiting regions outside Soviet boundaries have 
not been given. Synonyms of a group above the genus have not been given 
and no references are cited for descriptions; these, unless otherwise stated, 
are from Simpson (1945). 

The names given to domestic forms, as adopted and clearly explained 
in Volume I, have not been used for wild species; the name given to the 
wild form has been used as the species name. 

The total number of species in the class is about 3,500^ and the 
number of specie in Russian fauna about 300 (Heptner, 1956). In fact, 
recently an intense study of karyotypes in some cases has led to modifica- 
tions of apparently natural patterns, which were ascertained in the last decade 
by the morphogeographic method. Studies based on karyosystematics often 
result in the recognition of an increased number of species. However, at 
present, only a small number of species of this class have been studied 
from this viewpoint, mostly isolates of various species. It is therefore still 
difficult to establish definitely the extent of such an increase. Judging from 
the fact that whatever has been described is based on special studies of 
Russian fauna, major revisions relating to the number of species in the class 
are not likely. 

In the present volume A.A. Sludskii compiled the sections "Biology" 
and "Practical Significance" for all species except the lion. The remaining 
text has been written by V.G. Heptner, who also compiled all the range 
maps except those for which compilers have been duly acknowledged. V.G. 
Heptner is responsible for the general classification, limits, number, and 
sequence of species; number, limits, and identification of subspecies; and 
all the synonyms adopted in this volume. 

Almost the entire literature published up to 1967 and some of 1968 
has been reviewed. Only in a few cases are reports from more recent works 
occasionally cited. Information on the Amur cat, the European forest and 
steppe cats and the biology of hyaena has been greatly supplemented or 
elaborated from the latest information of early 1971. Only cited works have 
been referenced in the Literature Cited. References to literature have 
generally been placed in parentheses in the text, giving the surname of the 
author (without initials) and year of publication; initials are given only in 
those cases when more than one scientist has the same surname. References 

2E. Mayr (1969) estimates 3,700 and Th. Haltenorth (1%9) 4,146. 


to persons who have provided verbal or written communications of 
unpublished data are identified, unlike published references, without a date 
but with the initials of the author. 

Naturally, both authors of the present volume have extensively 
incorporated in the text their own unpublished data, especially since cats 
have long attracted their attention. In addition, various types of informa- 
tion still unpublished have been kindly fumished by V.K. Abramov 
(Vladivostok), Yu.V. Averin and M.N. Lozan (Kishinev), N.K. Vereshchagin 
(Leningrad), G.D. Dul'keit (Stolba preserve, Krasnoyarsk), N.I. Ishadov 
(Ashkhabad), B.A. Kuznetsov (Moscow), O.N. Nur-Gel'dyev (Ashkhabad), 
V.E. Prisyazhnyuk (Sudzukhin preserve), Yu.F. Sapozhenkov (Kostroma), 
V.N. Skalon (Irkutsk), N.D. Sysoev (Vladimir), B.F. Tserevitinov (Moscow), 
K.K. Chapskii (Leningrad), and many other colleagues, including workers 
in several regional game farms, who have been duly acknowledged at the 
appropriate place in the text. G.F. Bromlei and A.G. Pankrat'ev have 
provided particularly important data on the Far East. Pankrat'ev also 
furnished extremely valuable systematic collections. 

Sketches of animals in both color and black and white, as in the 
preceding volumes, are by the famous wildlife painter A.N. Komarov. 
Drawings of skulls and all anatomical sketches are by the zoologist-artist, 
N.N. Kondakov. They have prepared all the foregoing especially for this 
book from specimens in the Zoological Museum of Moscow University. 
Only the drawing of the hyaena skull was made by V.N. Lyakhov. The 
photographs in the biological sections of this work have been taken in part 
from other works by V.G. Heptner. Photographs have also been provided 
by V.K. Abramov (Vladivostok), Yu.K. Gorelov (Badkhyz preserve), S.V. 
Marakov (Kirov), AG. Pankrat'ev (Vladivostok), V.E. Prisyazhnyuk 
(Sudzukhin preserve), and others, whose names have been duly 
acknowledged at the appropriate place. All photographs, with a few 
exceptions, are published here for the first time. Moreover, all illustrations 
are original and have been selected by V.G. Heptner. 

The constant cooperation of the Chief Librarian of the Zoological 
Museum of Moscow University, V.I. Korotkova, especially in making 
available rare publications, is deeply appreciated. 

Cordial acknowledgement is given here to all the persons listed above 
and to others who in any way assisted in the preparation of this work. The 
authors are particulariy grateful to Prof. I.B. Volchanetskii (AM. Gorkii 
Kharkov State University) and to Prof. A.P. Komeev (T.G. Shevchenko 
Kiev State University), who took upon themselves the task of reading the 
manuscript and offering many useful comments, and to our colleague, LP. 
Mitina of the Department of Vertebrate 2Loology, who provided extensive 
and competent editorial assistance. 

This work was mostly executed within the walls of the Zoological 


Museum of Moscow University and is based to a great extent on the rich 
collections held there.-' The authors are deeply obliged to the Director of 
the Museum, O.L. Rossolimo and to all the workers in the Division of 

The first two publications under this monograph have been well received 
in the Soviet Union and abroad. The German translation of Volume I (G. 
Fischer, Jena, 1966) has also been well reviewed by the international press. 
The authors hope that this volume likewise will enjoy a good reception. 
They request all those using this publication to communicate errors and 
suggestions for improvement to Prof. Vladimir Georgievich Heptner, 
Zoological Museum of Moscow University, 6 Gertsen Street, Moscow 
103 009. 

-^Museum material has been designated by the abbreviation ZMMU (or KZMMU) 
and that of the Institute of Zoology of tlie Academy of Sciences of the USSR as ZMAN 
(or KZMAN). 

Table of Contents* 






Key for Identification of Families in the Order Camivora 1 

Family HYAENIDAE** Gray, 1869 (Hyaena) 2 

Subfamily Hyaeninae** Mivart, 1882 8 

Genus Hyaena Brisson, 1762 (Striped Hyaena) 8 

Hyaena hyaena Linnaeus, 1758 10 

Diagnosis 1 1 

Description 1 1 

Systematic Position 16 

Geographic Distribution 16 

Geographic Variation 20 

Biology 21 

Practical Significance 45 

Family FELIDAE Gray, 1821 (Cats) 47 

Key for Identification of Species of the Cat Family, Felidae 66 

Genus Panther a Oken, 1816 76 

Panthera leo Linnaeus, 1758 (Lion) 83 

Description 83 

Systematic Position 86 

Geographic Distribution 87 

Geographic Variation 92 

Biology 94 

*Pages 550-551 in Russian original. The English table of contents is not a literal 
translation — Sci. Ed. 

**Transposed in Russian original — Sci. Ed. 


Panther a tigris Linnaeus, 1758 (Tiger) 




Systematic Position 


Geographic Distribution 


Geographic Variation 




Practical Significance 
Panther a pardus Linnaeus, 1758 (Leopard) 





Systematic Position 


Geographic Distribution 


Geographic Variation 




Practical Significance 


Genus Uncia Gray, 1854 (Snow Leopard) 


Unci a uncia Schreber, 1775 




Systematic Position 


Geographic Distribution 


Geographic Variation 



Practical Significance 
Genus Felis Linnaeus, 1758 (Small Cats) 


Felis (Felis) euptilura Elliot, 1871 
(Amur, or Far-eastern Forest Cat) 




Systematic Position 


Geographic Distribution 


Geographic Variation 


Practical Significance 



Felis (Felis) chaus Giildenstaedt, 1776 
(Jungle Cat) 






Systematic Position 


Geographic Distribution 


Geographic Variation 




Practical Significance 


Felis {Felis) silvestris Schreber, 1777 
(Wild Cat) 




A. Group silvestris (European 
Forest Wildcats) 




Systematic Position 


Hybridization of Wild with Domestic Cats, 
and Hybrid Populations 


Geographic Distribution 


Geographic Variation 




Practical Significance 


B. Group libyca (Steppe Wildcats) 




Origin of Domestic Cats 


Systematic Position 


Geographic Distribution 


Geographic Variation 




Practical Significance 


C. Felis (Lyfix) caracal Schreber, 1776 






Systematic Position 



Geographic Distribution 


Geographic Variation 




Practical Significance 


Felis (Lynx) lynx Linnaeus, 1758 (Lynx) 






Systematic Position 


Geographic Distribution 


Geographic Variation 




Practical Significance 


Felis (Otocolobus) margarita Loche, 1858 
(Sand Cat) 






Systematic Position 


Geographic Distribution 


Geographic Variation 




Practical Significance 


Felis {Otocolobus) manul Pallas, 
1776 (Manul) 






Systematic Position 


Geographic Distribution 


Geographic Variation 




Practical Significance 


Genus Acinonyx Brookes, 1828 


Acinonyx jubatus Schreber, 1775 (Cheetah) 






Systematic Position 



Geographic Distribution 709 

Geographic Variation 714 

Biology 717 

Practical Significance 727 



*The Russian original also included an Index of Russian Names, omitted here. 

Systematics of the Class Mammalia 

The classification of major subdivisions (orders and higher taxa) adopted 
in the present publication is based on the systematic monograph of the class 
by Simpson (1945), who extensively relied on paleontological data. It is 
very close to the system of classification proposed by M. Weber (1928), 
which in its time was well founded on the anatomy of extant and fossil 
forms. The orders proposed by these zoologists have long been widely 
recognized and their grouping into higher taxa as suggested by Simpson, 
raises no serious objections. 

Extreme division of the orders is a recent trend. Thus marsupials have 
been divided into three and even five independent orders (opossums — 
Didelphida, marsupial martens — Dasyuria, marsupial badgers — Peramelia, 
caenolestids — Caenolestia, and marsupials, wombats, and kangaroos — 
Phalangeria), cetaceans into two (toothless whales — Mystacoceti and toothed 
whales — Odontoceti), primates into three (lemurs — Prosimiae, simians — 
Simiae, and tarsiers — Tarsioidea), and so on. All these proposals, however, 
are not yet well founded. Moreover, among mammalian systematists extreme 
division of the orders is not generally favored. Neither is it commended 
here. An order should be primarily an integrating and not a divisive concept. 
Otherwise the same situation will arise in mammalogy as has already 
occurred in ornithology, where the concept of an order in some systems 
of classification has largely lost its significance. As a result the scientific 
aspect of systematics of vertebrates has been undermined. 

In the classification of orders under consideration here, only one change 
has been made, i.e., pinnipeds have been elevated to an independent order; 
usually they are classed as a suborder of Carnivora. Judging from the 
systematics of recent mammals, these two groups have diverged and followed 
different directions of specialization. The isolation of pinnipeds in a separate 
order is as justified as the isolation of some other orders. 

Based on already established views, Lagomorpha are also treated here 
as an independent order (Weber considers them a suborder of rodents). Hence 
instead of the 17 orders of Weber and the 18 orders of Simpson, we propose 
19. Of these, 10 (56.6%) are represented in our fauna, one of them — sirenians 
is now extinct (Steller's sea cow; see Volume II, Part 1). 

The main subdivisions within the orders in most cases adhere to 
Simpson's scheme. By and large his classification is the most well founded. 
Nevertheless, significant deviations do occur in the present work, mainly 


at the level of family and especially at the level of genus. Our concept of 
genus is much broader than Simpson's. In our opinion a broader concept 
of generic divisions conforms better to the structure and principles of natural 
classification. The classification of cats has consequently undergone a general 
revision in this book. 

The systematic classification is given in the chart below. Orders present 
in our fauna are marked with an asterisk. (V.H.) 



Subclass THERIA 

Infraclass METATHERIA 
Infraclass EUTHERIA 


Cohort GLIRES 

Cohort MUTICA 





















Key for Identification of Orders^ 

1 ( 2). Hind limbs absent; fishlike body with large bilobate caudal fins 

set horizontally CETACEA. 

2 ( 1). Hind limbs present; body not fishlike and tail, if present, not in 

the form of a bilobate fin. 

3 ( 4), Forelimbs in the form of leathery wings CHIROPTERA. 

4 ( 3). Forelimbs of a different structure. 

5 ( 6). Fore- and hind limbs very short and resemble paddles, i.e., all 

the digits entirely enclosed in a common integument cover . . . 

6 ( 5). Fore- and hind limbs of a different structure, not paddle-shaped. 

7 (10). Feet possess hooves. 

8 ( 9). Only onet hoof on each limb *PERISSODACTYLA. 

9 ( 8). On each limb, two large hooves and two small ones above them 


10 ( 7). Hooves absent (claws present). 

11 (14). Between large chisel-shaped incisors and molars on jaws, a large 

toothless gap (diastema) occurs; its length not less than length 
of entire row of molars on corresponding jaw. Canines absent. 

12 (13). Two incisors on upper jaw RODENTIA. 

13 (12). Four incisors on upper jaw; behind each large shaф incisor is 

located a minute blunt tooth LAGOMORPHA. 

14 (11). Diastema between incisors and molai^s absent or much smaller 

than length of molar row. Canines present. 

15 (16). Anterior portion of muzzle extended into well -developed small 

proboscis. Anteriormost tooth on each jaw, or only upper jaw, 
much larger than adjacent tooth^ INSECTIVORA 

Orders marked with a single asterisk have been described in Volume I and those with 
two asterisks partly in Volume II and partly in Volume I.* 

*No order was double-asterisked in the Russian text — General Editor. 

^This refere only to horses, the only perissodactyls in the Soviet fauna. It may be noted 
that many perissodactyls have more than one toe on their legs. In fact, their distinctive feature 
is the odd number of toes — General Editor. 

^Limbs differ in structure in camels. The leg ends in a broadened callu-sed padding 
and true hooves are absent. There are tss'o highly broadened nails. Camels have not been 
included in this table as they are domesticated animals (wild camels are extinct). 

^If the structure of teeth is different, proboscis is always present and furthermore the 
front leg extremely short, feet very broad with huge claws, and set on edge with inner surface 
facing backward (moles). 


16 (15). Anterior portion of muzzle does not form a proboscis. Anteriormost 

tooth on each jaw not larger than one next to it 


■^On the coast of the Commander Islands, skulls of the extinct Steller's cow — a 
member of the order of sea cows (Sirenia) — have been found. They are distinguished by 
upper and lower jaws measuring 60 cm in length, in which teeth or traces of them (alveoli) 
are absent. Steller's cow has been described in Volume II, Part 1. 

Superorder FERAE Linnaeus, 1758 

Order CARNIVORA Bowdich, 1821 ^ 

Key for Identification of Families in the Order 

Identification by External Characters 

15 1 ( 6). Five toes on hind feet. 

2 ( 3). Tail short (shorter than hind foot) and hidden in body fur or barely 

protrudes. Size very large; body length of adults 1.4 to 2.0 m or 
more bears, family Ui'sidae. 

3 ( 2). Tail long (longer than hind foot) and invariably protmdes distinctly 

from fur. Body length less than 1.2 m. 

4 ( 5). Tail distinctly bicolored; six or seven dark-colored transverse rings 

displayed on lighter background 

raccoon, family Procyonidae. 

5 ( 4). Tail monochromatic or terminal part often darker than base, but 

transverse dark-colored rings absent 

weasels, family Mustelidae. 

6 ( 1). Four toes on hind feet. 

7 ( 8). Four toes on front feet. Neck and back with tall shaggy mane 

of long resilient hair. Color gray (without rusty tinge) with black 
stripes traversing trunk hyaena, family Hyaenidae (p. 2). 

8 ( 7). Five toes on front feet (first toe set above rest). Tall mane along 

neck and back absent. Coloration different. When trunk covered 
with black transverse stripes, bright rust-colored tinge of body 
very vivid. 

9 (10). Head elongate, with long muzzle similar to that of a dog or fox. 

Tail always reaches tarsal joint and is furry, but hair at base short 
and thinner than in the middle. Claws nonretractile; tips set forward 
and downward, not highly compressed laterally, and relatively 

16 blunt. Coloration without vivid spots or transverse stripes. . . . 
wolves, etc., family Canldae. 

^Characteristics of tlie order have been given in Volume II, Part 1. 
^Families for which the page number is not shown have been described in Volume II, 
Part 1. 

10 ( 9). Headround, with short muzzle like that of a cat. Tail reaches tarsal 
joint or is shorter; uniformly covered with hair throughout its 
length, not thinner at base than in the middle. Claws retractile; 
tips set forward and upward [when retracted], highly compressed 
laterally, and very shaф. If claws nonretractile, then body size large 
(like that of a large dog) and color of body light, with small, shaфly 
defined but not circular black spots. Coat with spots, or transverse 
stripes, or uniform in color cats, family Felidae (p. 47). 

Identification by Skull Characters 

1 ( 6). Palatines extend back beyond transverse line drawn across posterior 

margin of last teeth by less than one-half their width between those 

2 ( 3). Four teeth [or three] on each side of lower jaw behind canine; five 

teeth in upper jaw. Total number of teeth 34 or 32* 

hyaena, family Hyaenidae (p. 2). 

3 ( 2). Dental formula different. 

4 ( 5). Three teeth on each side of lower jaw behind canine; four or tliree 

in upper jaw. Total number of teeth 28** or 30 

cats, family Felidae (p. 47). 

5 ( 4). Seven or six teeth behind canine in lower jaw and six in upper 

jaw. Total number of teeth 42 or 40 

wolves, etc., family Canidae. 

6 ( 1). Palatines extend back beyond transverse line drawn across posterior 

margin of last teeth by more than one-half their width between 
those teeth.-^ 

7 ( 8). Skull large, maximum length exceeds 200 mm 

bears, family Ursidae. 

8 ( 7). Skull small; maximum length less than 200 mm. 

9 (10). Six teeth behind canine in upper jaw and six in lower jaw. Total 

number of teeth 40 raccoon, family Procyonidae. 

10 ( 9). Less than six teeth behind canine in upper jaw and four, five, 

or six in lower jaw. Total number of teeth 38 or less 

weasels, family Mustelidae. (V.H.). 

Superfamily FELOIDEA Simpson, 1931 

Family HYAENIDAE Gray, 1869 

Hyaenas are specialized, often highly specialized, carnivores of large size. 

* Error in Russian original; reads "24" — Sci. Ed. 

**Error in Russian original; reads "38" — Sci. Ed. 

■^Measurement should be taken along the suture between the palatines. 

Trunk relatively short, fairly massive, on the whole wolf-like, but rear 
portion weak and low, withers high, and line of spine slopes down noticeably 
toward croup. Forelegs high, but hind legs very short. Neck quite short 
and thick. Head generally resembles a wolf's, but much larger, massive, 
and heavier, with a very powerful, relatively short facial portion and heavy 
lower jaw. 

Forelegs tall and thin, somewhat bent. Hind legs also appear thin and 
relatively weak, particularly the thigh. Limbs typically digitigrade; fore- and 
17 hind paws with four digits each (digit I absent; in one species five digits 
occur on forepaws). Digits III and IV equal in length, but shorter on average 
than either digits II or V. Digits with short, powerful, straight, blunt, and 
nonretractile claws. Digital pads highly developed; digits connected by a 
dense, thick, and elastic membrane which reaches pad. At base of digits 
a large callused cushion, roughly semi-crescentoid in shape, occurs. Small 
carpal pad present high amidst fur on forefoot; pad absent on hind foot. 
Digital pads bulge notably. Digits always set close to each other (ball-like). 

Pelage sparse and coarse, with underfur poorly developed or lacking; 
majority of species have rich mane of long hair running along spine from 
withers or from head. Color dull gray or brown or rusty-brown, more or 
less uniform, or with dark (black) transverse stripes or spots. 

Perineal scent glands absent, but tract (pocket) in anal region with well- 
developed anal and large sebaceous glands. 

Skull massive and heavy with powerful facial portion perceptibly 
extended forward, and compressed, and very small brain case. Viewed from 
the top, skull appears wedge-shaped with very broad (especially from the 
rear) and powerful zygomatic arches diverging at an angle. As a result of 
very powerfully developed sagittal crest, line of dorsal profile from 
interorbital region to occipital is greatly elevated. Occipital region also highly 
compressed; when skull viewed from the rear, it has a triangular shape. 
Sagittal crest forms the extended shaф apex of this triangle. Broadly 
developed zygomatic arches and compression of brain case associated with 
development of very powerful masticatory muscles. 

Alisphenoid canal well developed or absent. External auditory meatus 
in the form of a tube. Auditory bulla swollen; externally its division into 
two parts not perceptible; internally, partition wall shifted far back and 
entotympanic portion very small. Paroccipital processes large and massive, 
projecting down beyond lower margin of tympanic bulla. Posterior palatine 
foramina shifted far anterior, to level of second premolar. 

Dental formula: 

1 , с , pm , m = 34 or m = 32. 

3 13 1 1 

In several respects, dentition is similar to that of the cat family (Felidae). 
In most species it is very powerful, indeed one of the most powerful in 
the order, and highly specialized for handling coarse food and crushing 
bones. Carnassial teeth (upper fourth premolar and first lower molar), 
especially the upper, very powerful and shifted far posterior to the point 
of exertion of utmost pressure on the jaws. Inner cusp (protocone) of upper 
carnassial tooth well developed, shifted to inner surface of anterior end of 
tooth, and set at a right angle. 

Other teeth, apart from first upper premolar, and nonfunctional or almost 
nonfunctional and very poorly developed (rudimentary) upper molar, very 
powerful with broad bases and cutting edges, but not very sharp cusps. 
Canines powerful and thick, but not long. As a result of feeding on insects, 
one genus (Proteles) has highly deviant teeth (see below) and no upper 
molar whatsoever; this tooth is sometimes absent in the spotted hyaena 
(Crocuta) also. 

Vertebral column, including cervical region, of limited mobility. Os 
penis not developed. 

Most species are specialized carrion eaters, mostly of ungulates, and 
sometimes of marine strandings; one species is specialized for feeding on 
termites and ants. More rarely, they hunt live animals, but mostly prey on 

18 the sick and wounded or the young and helpless. They are nocturnal, mostly 
solitary animals. The den is generally concealed in burrows (mostly those 
of porcupines), rock crevices, caves, and so on. 

The geographic range (Fig. 1) is associated with relatively arid regions 
of the Old Worid. It occupies all of Africa, from the Cape of Good Hope 
to the Mediterranean Sea, except for the desert regions of North Africa 
(Libyan desert and others) and the region of dense humid tropical forests 
(hyaenas are known, however, in Angola, Togo, Cameroon, and Congo), 
Arabian peninsula, Asia Minor, Iraq, Iran, Trans-Caucasus, Afghanistan, 
southern part of Middle Asia, India from the Himalayas to Cape Comorin 
[Kanyakumari] and in the east as far as Bengal and Nepal-Terai (absent 
in Burma and Assam). All trae hyaenas (striped and spotted — genera Hyaena 
and Crocuta) are associated in their distribution and attain maximum 
population levels in regions where ungulates are numerous. 

The family as a whole represents in all respects a very distinct group, 
the independence of which is unquestionable. In spite of similarity with 
wolves in appearance, the affiliation of the family to the superfamily of 
cats is also clear. The hyaena family occupies a somewhat intermediate 

19 position between the relatively generalized family of civets (Viverridae) 
and the extremely specialized family of cats proper (Felidae). In some 
respects, hyaenas are closer to the latter. 

Some species of the family, true hyaenas, are a monotypic and compact 


Fig. 1. Reconstructed geographic range of the family Hyaenidae. V.G. Heptner. 

group. However, one genus (Proteles) deviates very sharply. Although a 
typical representative of the family in its main features, it has several unique 
features. For example, its dentition is degenerate, associated with its diet 
of ants and termites: all the teeth are very small, weak, and wide-set; the 
upper and lower rows together do not form a single cutting apparatus; and 
most of the teeth do not come into contact with each other when the jaws 
are closed. The canines and incisors are also as weak as the cheek teeth. 
Upper molars are absent aUogether and the teeth are easily lost during life. 
Such a dentition cannot serve as an effective "defensive weapon"; thus 
their ability to spray a malodorous liquid from a preanal pouch provides 
supplementary protection. 

These characters are important and clearly separate the genus Proteles 
from true hyaenas {Hyaena, Crocuta). Some systematists regard these features 
as a justification for placing this genus in a separate family (Protelidae), 
adjacent to the hyaenas proper — Hyaenidae. The view has also been expressed 
that Proteles is essentially very closely related, or even belongs to the civet 
family (Viverridae), constituting a special subfamily within this family. From 
this viewpoint the similarity of Proteles to the hyaenas, Hyaenidae, is no 
more than convergence. Such a view has not been accepted, however. 

It would be proper to regard Proteles as a specialized form of hyaena, 
evidently separated long ago from the main line of development and hence 
preserving some characters of viverrids, the family from which hyaenas 
originated. However, the main thing is that all features of the aardwolf are 
essentially degenerate features of true hyaenas; there are no major qualitative 
differences. Hence, systematists of morphological emphasis do not separate 
the aardwolf into a special family; it is generally allocated to a distinct 
subfamily. Thus the hyaena family consists of two subfamilies — true 
hyaenas, Hyaeninae, and the aardwolf, Protelinae. 

The skull features, especially dentition of the aardwolf as already pointed 
out, are associated with their specialization for feeding on insects (termites 
and ants). This feature is also responsible for some other characters, for 
example, their low mobility. This is one of the very rare, if not unique, 
instances of extreme specialization of a large carnivore for feeding on insects. 

In origin, the family is distinctly linked with the viverrids (Viverridae) 
and may be considered descendant from it. The whole family is relatively 
young and became distinguishable in the Miocene. The most primitive are 
the Ictitheriinae, a unique subfamily known from the Lower and Middle 
Miocene of western Europe to eastern Asia. These are viverrid-like 
carnivores, usually considered the group ancestral to true hyaenas 
(Hyaeninae) and the aardwolf (Protelinae). Some, such as Progenetta, are 
sometimes placed with the Viverridae. However, the Ictitheriinae represent, 
evidently, only a lateral dying branch of the hyaena family. The aardwolf, 
Protelinae, among extant animals, is closest to the Ictitheriinae. 

The remaining fossil forms belong to contemporary groups; all three 
lineages — spotted hyaena, striped hyaena, and aardwolf — diverged quite 
early, i.e., in the Miocene. Spotted hyaenas (Crocuta) were particularly 
abundant in the Old World. In Europe this group became extinct only in 
the Pleistocene. Here it was depicted toward the end of its existence as 
the well-known "cave'" hyaena, C. spelea. The genus Hyaena appeared 
in the Upper or Middle Pliocene, while the genus Lycyaena (representing 
the ancestral group) lived in the Lower Pliocene. Throughout its history 
the family was confined to the Old World and only one species is known 
from the Lower Pleistocene of North America. 


20 The family is very depauperate. In contemporary faunas, there are only 
three genera and four species: aardwolf, Proteles Geoffroy, 1824 with one 
species {P. cristatus Sparrm., 1883); spotted hyaena, Crocuta Каир, 1828 
with one species (C. crocuta Erxl., 1777); and striped hyaena. Hyaena Briss., 
1762 with two species (striped hyaena, H. hyaena Linn., 1758 and brown 
or shore hyaena, H. brunnea Thunb., 1820), 

Of the three genera, two (Proteles, Crocuta) wholly belong in Africa 
and the area south of the Sahara (Ethiopian region), and one genus (Hyaena) 
with two species (see below) is distributed in Africa (Palearctic and Ethiopian 
regions), the Near East, and southern Asia. 

The economic importance of this group is possibly more favorable 
(scavenging of carrion) than unfavorable, although this differs in some places 
in accordance with population density, nature of cultivation, etc. Sometimes 
(southern Africa), hyaenas cause damage to animal husbandry (by killing 
sheep) and even pose a threat to man. 

ТЪеге is only one genus, Hyaena Briss., 1762, found in the Soviet Union. 

Subfamily Hyaeninae Mivart, 1882 

21 Genus of Striped Hyaenas 

Genus Hyaena Brisson, 1762 

1762. Hyaena Brisson. Regn.Anim., 2nd ed., pp. 13 and 168. Canis hyaena 

Linnaeus, 1758. 
1868. Euhyaena Falconer. Paleont. Memoires, vol. 2, p. 464. Canis hyaena 

Linnaeus, 1758. (V.H.) 
In general appearance and form the body is wholly typical of the subfamily. 
Ears large, tall, and pointed. Hair cover relatively long with a large mane 
of very long, erect, coarse hair along the spine. Color gray with transverse 
dark stripes on trunk. Newborn with same color pattern as adults. External 
genitalia of females normal.^ 

Upper molar small but always present and relatively well developed; 
its transverse diameter is approximately three times greater than its length. 
It is shifted lingual to the tooth row and set at a right angle to the longi- 
tudinal axis of the camassial tooth at the level of its posterior margin (as in 
various species of the cat family, Felidae). The lower carnassial tooth has 
two large anterior cusps externally [buccal] and a small one internally 
[lingual], away from the second outer cusp. On the tooth a small but well 
developed heel occurs, with a tiny tubercle set opposite the upper molar. 
Unlike the upper jaw, the third and fourth premolars and molar of the lower 
jaw are almost identical in size, strength, and massive and relatively broad; 
the camassial (molar) is equal to the fourth premolar or only somewhat 
larger than it. All the teeth together form a large and powerful tearing and 
crushing instrument, occupying much of the dental row, in the region of 
maximum exertion of force. The outer upper incisors are twice as large 
as the inner ones. 

Striped hyaenas are solitary animals, specialized for feeding on carrion 
(mainly ungulates) and occasionally on marine organisms washed ashore. 

The reconstructed range of the genus generally coincides with the range 
of the family. At present striped hyaenas (brown, H. brunnea) are extinct 
in the southernmost region of Africa. The situation is evidently similar in 

4n the genus Crocuta (spotted hyaena) the structure of the external genitaha of females 
is altogether different and resembles that of males. The characteristic features of the genus 
Hyaena have been compared with those of the genus Crocuta. 

some other parts of the mainland, particularly in the northern edge of the 
range of the brown hyaena in eastern Africa. 
22 The taxonomy of the genus (not taking into consideration the geographic 

variation of the species) is generally fairly well known; the genus consists 
of just two species: the numerous so-called species of striped hyaena 
described from Africa and Asia, especially by Matschie and Satunin, have 
the status of neither species nor race. 

Of the two genera forming the subfamily of true hyaenas (Hyaeninae), 
the genus Hyaena should be considered somewhat less specialized than the 

Fig. 2. Reconstructed species range of striped hyaena, Hyaena hyaena Linn. V.G. 



genus CrocLita (spotted hyaena). This is supported particularly by the 
frequent absence of the upper molar, the general strengthening of the hyaena- 
type skull, its very large size, the atypical structure of external genitalia 
(formerly interpreted as attributable to hermaphroditism), the spotted 
coloration, variation in coloration of newborns and adults, and other features 
of the spotted hyaena. 

The genus Hyaena is known from the Middle Pliocene of Europe 
(Moldavia and western Europe) and from the Pleistocene. Hyaenictis from 
the Lower Pliocene of Europe and Lycyaena from the I^wer and Middle 
Pliocene of Europe and Asia (China; regarded usually as a predecessor of 
the former) are closely related to it. The genus Crocuta had already evolved 
in the Upper Miocene of Asia whence, as also in Europe, it is known until 
the Pleistocene. Both of these groups of hyaena have survived as independent 
branches for a very long time (from the Miocene). 
23 There are only two species in the genus: the brown or shore hyaena, 

H. brunnea Thunberg, 1820, and the striped hyaena, H. hyaena Linnaeus, 
1758. Both species are found in Africa. Brown hyaenas are distributed south 
of the Sahara, as far as the extreme south of the continent (reconstnicted 
range), except for regions of dense forests in western Africa; striped hyaenas 
occupy the northern region of Africa and penetrate south of the Sahara, 
mostly in the eastern half of the mainland (see "Description" of species). 
There is only one species in the Near East, Middle Asia, and southern Asia 

One species, the striped hyaena {Hyaena hyaefia Linnaeus, 1758) lives 
in the Soviet Union and represents about 0.3% of species of the fauna. 

The range covers the extreme south of the Soviet Union, namely, the 
Trans-Caucasus and the southernmost regions of the western half of Middle 

Economically speaking, partly because of its small numbers, the hyaena 
has no impact, but is useful in scavenging carrion. Some damage to cattle 
is not ruled out. Sometimes, apparently, this animal poses a danger even 
for man. (V.H.) 


Hyaena hyaena Linnaeus, 1758 

1758. Canis hyaena. Linnaeus. Systema Naturae, 10th ed., vol. 1, p. 40. 

Benna mountains, Laristan, southern Iran (not India as stated by 

Ognev, 1931; and others). 
1777. Hyaena striata. Zimmermann. Spec. Zool. Geogr., p. 366. Renaming 

of hyaena Linnaeus. 


1820. Hyaena fasciata. Thunberg. Sv. Vet. Akad. Hand!., vol. 1, p. 59. 

Renaming of hyaena Linnaeus. 
1844. Hyaena vulgaris indica. Blainville. Osteogr. Mamm., vol. 2, pi. 6. 

1905. Hyaena vulgaris zarudnyi. Satunin. Izvestiya Kavk. Muzeya, vol. 2, 

p. 7. Lower course of Kamna River, southeast Iran. 
1905. Hyaena bokharensis. Satunin. Ibid., vol. 2, p. 8. "Eastern Bukhara" 

(southern Tadzhikistan). 
1905. Hyaena bilkiewiczi. Satunin. Ibid., vol. 2, p. 9. Ashkhabad region. 
1905. Hyaena vulgaris satunini. Matschie. Sitzungsber. Ges. Naturf. 

Freunde Berlin, p. 363. Trans-Caucasus. (V.H.) 


H. hyaena L. is the only species of the genus inhabiting the Soviet Union. 


At first sight, the hyaena seems to resemble a wolf or large dog, but possesses 
an extremely typical external appearance (Fig. 3). Trunk fairly massive but 
short and set on long legs. Forelegs significantly longer than hind ones, as 
a result of which the body is raised at the withers, but the croup significantly 
lower, and the back slopes down. The high withers are the result not only 
of the difference in leg length, but also intensity of growth of the spinous 
processes of the corresponding vertebrae, and of the mane running along 
the back. Legs relatively thin and weak; forelegs perceptibly bent in сафа! 
region. Head placed on thick and fairly long but poorly mobile neck; heavy 
and massive, with a small, noticeably blunt facial portion. Eyes rather small. 
Ears very large, broad, set high, and pointed at the tip; wide-set, directed 
shaфly upward (do not "droop" sideways) and incline somewhat forward. 
Paws have bulky pads, characteristic of all hyaenas, and powerful but short, 
24 blunt claws, which appear trimmed off at the tips. Tail not long and terminal 
hairs do not descend below calcaneal joint. 

For an animal of this size, the winter coat is quite long and fairly 
uniform, with narrow but luxuriant mane of long resilient hair along 
back from occiput to base of tail. Mane usually "falling" but, on occasion, 
hairs are capable of standing more or less vertically when erected. Tail 
also covered with long hair, which appears shaggy. Coat generally coarse, 
in places bristly, and varies greatly in different seasons. In winter, coat 
fairly dense, relatively soft, with well-developed underfur and long hair. 
Guard hairs on trank (sides) are at this time 50 to 75 mm long, mane 
hairs 150 to 225 mm, and tail hair about 150 mm. The summer coat is 


Fig. 3. Striped hyaena, Hyaena hyaena Linn. Sketch by A.N. Komarov. 

very coarsCj much shorter than in winter, sparse, and underfur is absent; 
the mane is nonetheless large. 

General shade of winter coat usually dirty brownish-gray or dirty gray. 
Hair of the mane wide, with pure black or light brown hair terminally and 
nearly as wide with light gray or almost white hair at base. Small dark bands 
alternate with much lighter colored sections in basal half of mane; as many 
as four such bands may occur (Fig. 4). 

Muzzle, up to eyes and chin, dark, grayish-brown, brownish-gray or 
black, while top of head and cheeks are lighter in color, usually dirty or 
grayish-brown. Ears very dark-colored, almost black. Inside as well as outside 
of ears covered with sparse fur, the color of which depends on skin color. 
Along edges of ears, hair somewhat more dense and white in color. On throat 
and downward from front of neck, large black spot separated from chin by a 
light zone. From flanks, dark field ascends to rear of cheek; some separate 


black spots occur below dark field. Similar dark spots scattered directly in 
25 front of shoulder; individual spots and short dark band located in mane 
on withers, and fairly long band extends forward and downward from mane 
through shoulders. 

Inner and outer surface of forelegs, almost as far as carpus, covered 
with small dark spots and transverse stripes. No dark patch on tail tuft; 
tuft usually lighter in color than tail itself. Four indistinct dark vertical stripes, 
or rows of diffused spots, or a combination of both, seen on flanks. Outer 
surface of thigh with three or four distinct vertical or oblique dark bands, 
merging into transverse stripes in lower portion of legs (to heel). Only spots 
present on legs below heel. Dark stripes and spots brownish-black. Belly 
dirty gray with a small number of pale spots; latter also present on inner 
side of thighs. Hair on upper part of tail with same color pattern as mane 
hair, i.e., black tips but no transverse dark-colored zones in light-colored 
part at base. Tip of tail black but underfur white. 

The coloration described above is subject to considerable individual 
variation with respect to general background, intensity of dark markings, 
often number and disposition of spots, distinct separation of bands, and 
other features. The general background varies from fairly dark to rather 
brown, or straw-colored, or even almost dirty white. Pelage color changes 

Fig. 4. Striped hyaena in winter coat. Badkliyz preserve, southern Turkmenia. 
December, 1961. Photograph by Yu.K. Gorelov. 


greatly depending on wear and fading; obviously, the color is brighter 
when the coat is new. Before and during molt the coat is especially dull 
and pale; at this time the dark shades in particular often fade to brown and 

Sexual dimoфhism in color is absent but seasonal color dimoфhism 
is noticeable. In the short summer coat the stripes and spots are a deeper 
black, the bands more distinct, and the color contrast far shaфer. 

Newborn young have a silky coat devoid of mane or long hair on the tail. 
The general basic color is white, muzzle gray, with a broad, dark band 
extending along the spine, and disappearing on the withers and sacrum. The 
tail is wholly white. From the shoulders to the sacrum, however, eight vertical 

26 black stripes are quite distinct; stray spots between these stripes are likewise 
well defined. A black stripe extends along the top of the neck (Indian animals; 
Pocock, 1941). 

A fairly large pouch without hair is located at the anal opening of males, 
and large anal glands open into it at some distance above the anus. Several 
large sebaceous glands also appear between the openings of the anal glands 
and above them. Scrotum small, covered with hairless skin, and situated 
in the bare region surrounding the anus. Penis long and preputial folds set 
far forward. External genitalia of females normal; vulva located in hairless 
preanal region, from which it is separated by a narrow strip of hair-covered 
skin. Large anal glands open into preanal area above anus (Pocock, 1941). 

27 There are three pairs of teats. 

The major skull features are wholly typical of the family and genus, 
in particular, development of very high sagittal crest; height of crest at the 
auditory bullae constitutes about one-half the condylobasal length of skull, 
as does height of nasal region (together with lower jaw). Muzzle short and 
broad, facial region almost equal to that of brain case or shorter. Postorbital 
processes well developed, and have rough surface. Frontal region between 
them inflated (Fig. 5). 

Anterior and lower margins of orbits bounded by ridge. Zygomatic 
arch with well-developed postorbital process, and together with supraorbital 
process of frontal forms posterior, upper, and lower borders of orbit. Orbit 
closed to a significantly greater extent than in other Russian carnivores 
(except cats), though not completely. Tip of anterior processes of frontal 
usually extends to posterior process of premaxilla. Anterior end of na.sals 
deeply notched. Tympanic bullae high (bulging), shaфly projecting 
downward, and coastricted anteriorly; their axes lie at an angle to each other. 
Palate long and broad; distance between posterior ends of carnassial teeth 
more than one-half length of palate. Deep depression (place of attachment 
of powerful muscles) occurs on angular processes of lower jaw. Paroccipital 
processes very large and massive. 


Fig. 5. Skull of striped hyaena, Hyaena hyaena Linn. Sketch by V.N. Lyakhov. 


Features of dentition have already been discussed under generic 

Measurements "in the flesh" of Russian hyaenas are not available. 
Indian hyaenas (five adult males and five adult females; from Pocock, 1941) 
are as follows: body length, males, 99.0-109.5 cm, females, 95.5-108.0 
cm; tail length, males, 27-34 cm, females, 27-30 cm; length of hind foot, 
males 20.0-21.5 cm, females, 19.0-22.5 cm; length of ear, males, 15.0-16.5 
cm, females, 13.7-15.8 cm; height at shoulders, male (1), 75 cm, female 
(1), 65 cm. 

Condylobasal length of skull in males (9), 210-220 mm*, females (9), 
202-220 mm*; zygomatic width, males, 149-164 mm, females, 147-155 
mm; interorbital width, males, 47-50 mm, females, 44-55 mm; postorbital 
width, males, 35-39 mm, females 35-40 mm (India; Pocock, 1941). 

Among hyaenas of Middle Asia (9) and Trans-Caucasus (1) without 
sex differentiation: condylobasal length, 204-224 mm; zygomatic width 
143.0-160.2 mm; interorbital width, 44.1-51.7 mm; postorbital width, 
32.2-39.9 mm; mastoid width, 74.1-90.2 mm (Ognev, 1931; material from 
Zoological Museum, Moscow University). Condylobasal length of an adult 
female from southern Turkmenia (Pul'-i-Khatum on the Tedzhen) was 221 
mm, zygomatic width 157.7 mm, interorbital width 45.9 mm, postorbital 
width 36.5 mm, and mastoid width 81.8 mm (V.G. Heptner). 

Males weigh about 38.5 kg, females about 34.0 kg (India; Pocock, 1941). 
There are apparently no significant sex-related size differences. (V.H.) 

Systematic Position 

The striped hyaena is closely related to other species of the genus, the African 
brown or shore hyaena, H. brunnea. The skull of the latter is larger and 
its dentition more powerful than in this species; because of these differences 
the brown hyaena may be considered a somewhat more specialized form. 
Its body hair coat is very long, the mane less distinct, color dark, and stripes 
less prominent. (V.H.) 

Geographic Distribution 

In open, mainly arid expanses of India, the Near East, and Africa except 
the southern half. 

Geographic Range in the Soviet Union 

The range (reconstructed) in the Soviet Union covers the extreme south 

"Error in Russian original: reads "cm" — Sci. Ed. 


of the country, in the Caucasus and Middle Asia, and consists of two separate 
sections — Caucasian and Middle Asian. These are continuous to the south 
28 outside the USSR. The northernmost edge of the range of the species as 
a whole is located in the Soviet Union (Fig. 6). 

In the Caucasus the range occupies the plains, foothills, and low 
mountains of the eastern Trans-Caucasus and displays a complex outline. 
It includes all of the montane part of Araks valley and the adjoining low 
hills (hyaenas have been sighted at Merg and Nakhichevan), in the Yerevan 
depression west of Echmiadzin, and as far as Surmala on the Araks, southwest 
of Echmiadzin. Hyaenas are distributed farther in the Mugansk steppe, 
Sal'yansk and Lenkoransk lowlands, and in the Talysh mountains, where 
the animals have been encountered at altitudes 2,000 m above sea level. 

In the north the range of the hyaena covers the area along the Alazan, 
Ion, and Kur', including Shirak (between Ion and Alazan) and Adzhinour 
(left bank of the Kur' below the mouth of the Alazan) steppes (Sheklinsk 
and Kartalinsk plateaus) and extends to the Karayaz forest (slightly east 
of Tbilisi) and almost to Tbilisi, and occupies the semideserts in the region 
of the right tributaries of the Kur' to the south of Akstafa (Idzhevansk and 
Shamshadinsk regions of Armenia). The northern boundary also encloses 
the foothills of the Great [Caucasus] range and extends from there southwest 
to the foothills and lower belt of the Karabakhsk range (Agdam and other 

Fig. 6. Reconstructed range of striped hyaena in the Soviet Union. V.G. Heptner. 


The Trans-Caucasian section of the range occupies the Apsheron 
Peniasula and extends in the form of a narrow projection along the coastal 
region of the Caspian Sea northward to the district between Derbent and 
Makhachkala — in any case to the lower reaches of the Samur or slightly 
farther north. Hyaenas extended as far as Akhta along the Samur. Mainly 
stray animals have been reported along the Caspian coast, possibly due 
to peculiar circumstances (military activities in the first half of the last 
century). A reported case of wandering as far as Ordzhonikidze (Vladikavkas) 
appears dubious. Reports of intrusions to Novorossiisk are undoubtedly 

In Middle Asia hyaenas have been sighted along the Atrek, evidently 
not reaching the sea, however; in the desert north of the river (Messeriansk 
desert), and throughout the Kopet-Dag, its foothills and temporarily to the 
adjacent northern plains, but never penetrating the highest parts of the 
mountains. In the Great Balkhan range, hyaenas are presently absent but 
may have lived there in the past. Definite information about this is absent, 
however. Information about occurrences along the Uzboi (Dement 'ev, 1955) 
is extremely dubious. To the east the range occupied the Tedzhen valley 
northward, evidently to the city of Tedzhen (possibly even lower), the 
29 Murgab valley, evidently to its lower reaches, all of the expanse between 
the upper divides separating the Murgab and Tedzhen (Badkhyz and 
Chengurets mountains), the Karabil district lying south between Murgab 
and Amu-Darya, and southeast Karakum at least to the Karakum canal in 
the north (villages of Zenkash, Neder-Belent, Kert-Kuyu, Aitysh-Kuyu, and 
others). Thus, in southeast Turkmenia hyaenas reached deep into the desert, 
albeit usually along their edges. 

Along the right bank of the Pyandzh and upper Amu-Darya, the range 
occupies a lairly significant section of the plciins, footliills, and low mountains 
from the Amu-Darya in the south to the foothills of the Gissar range in 
the north. In the east the range is bounded by the foothills of the Darvaz 
range and its southern spurs. The boundary evidently mns slightly east of 
70° or along it through the Kulyab, Muminabad, Bal'dzhuan, and Sarykhosor 
regions. From there, along the foothills of the Gissar range, it turned 
west, initially through the eastern part of the Gissar valley (region of 
Ordzhonikidzeabad), then through the Dushanbe region and B. Oktyabrask 
to Syry-Assiya in the upper course of the Surkhan-Darya, and probably even 
intersected the valley of this river slightly more to the north. From there 
the boundary encompassed the southeastern foothill spurs of the Gissar range 
and Kugitangtau and extended toward the Amu-Darya to the south of this 
mountain massif- — at Kelif. Precise data on temporary incursions of hyaenas 
along the Amu-Darya pertain to the Termez region and the lower reaches 
of the Shirabad-Darj'a. In the Amu-Darya valley, hyaenas have been reported 


from Chardzhou and Farab. These intrusions probably occurred from the 
south along the river valley. 

The natural range described above for Middle Asia remains more or less 
the same today. In the Caucasus, however, mainly before the twentieth 
century and more particularly in the last 30 to 40 years, it has fluctuated 
considerably; in fact, hyaenas almost disappeared completely from this area. 
Even in the 1930s they were encountered in small numbers only in some 
pooriy inhabited pockets of western Azerbaidzhan, eastem Georgia (then and 
later), the Kartalinsk plateau (in the region of the city of Udabno, southe<ist 
of Tbilisi), and probably Talysh. In the 1950s (F.F. Aliev) some stray animals 
(total of four) were caught in Azerbaidzhan at Yevlakhsk (on the Kur'), 
Agdzhabedinsk (Mil'sk steppe), and Gadrudsk (southern Karabakh range) 
region. In 1968 some hyaenas were encountered in the desert hills of 
Adzhinaur steppe (east of the lower Alazan), Kartalin and lori plateaus, 
the Araks valley within Nakhichevan Autonomoas Soviet Socialist Republic, 
and single individuals around Shemakha and in Karabakh (B. Agdamsk 
and Fuzulinsk regions). According to other data (Alekperov, 1966), hyaenas 
were absent in southwest Azerbaidzhan even in the early 1950s. Evidently 
this animal was very rare in the Shirak steppe (1969, Tsiteli-Tskara; A.B. 
Arabuli). The total disappearance of the hyaena from the Trans-Caucasus 
is apparently a recent phenomenon. Intrusions from Iran into the extreme 
south of the Trans-Caucasus can still be expected, however. 

Extensive hunting only partially explains the disappearance of the hyaena 
(in the Caucasus hyaenas are accused of carrying off children). More likely 
reasons would seem to be overall changes in weather conditions, reduction 
and often disappearance of ungulates, mainly goitered gazelle and in part 
large predators, and changes in the character of animal husbandry .^ 

According to some reports (Pidoplichko, 1951) contemporary species, 
or forms close to them, were widely distributed in the Quaternary Period 
in Europe. Remains have been reported from Ireland, England, Belgium, 
France, Spain, Portugal, Italy, Switzerland, Germany, Hungary, 
Czechoslovakia, and Poland, and in the USSR, in the Crimea, Ternopol, 
Dnepropetrov, and Odessa districts, around Kev, in Tatariya, the Kuban, 
and in the southern Urals. In Asia, outside the present-day range. Pleistocene 
30 hyaenas are known from the regions of Minusinsk, Krasnoyarsk, the Altai 
and Trans-Baikal regions, the Mongolian People's Republic, and China. 

However, most of these references evidently pertain to cave hyaena 

'Range based on works of Bogdanov, 1873; Dinnik, 1914; Satunin, 1915; Sultanov, 
1930; Ognev, 1931; Leviev, 1939; Heptner and Formozov, 1941; Vereshchagin, 1942, 1947, 
1959; Dal', 1954; Sekiinova et al., 1956; Heptner, 1956; Chernyshev, 1959; Nur-Gel'dyev, 
1960; Ishunin, 1961; Alekperov, 1966; and others; and unpublished data of V.G. Heptner 
and F.F. Aliev. 


(Crocuta spelaed), and fossils of the striped hyaena are known only from 
Paleolithic caves in Palestine, Morocco, Algeria, and Portugal. Pleistocene 
finds in France are less reliable; the direct association of the Pleistocene hyaena 
of England with present-day species is also dubious. According to these data 
(Vereshchagin, 1959), remains of striped hyaena have not been found 
anywhere in the Soviet plains, the Crimea, the Caucasus, and Siberia; fossils 
from the Paleolithic period have only been discovered south of Samarkand. 

Geographic Range outside the Soviet Union 

The range (reconstructed) in Asia occupies the Arabian peninsula, including 
its pre-Mediterranean part, Asia Minor, Iraq, Iran, Afghanistan (except the 
highest parts of the Hindu Kush), Baluchistan, and all of the northern Indian 
subcontinent to lower Kashmir, Nepal, and lower Bengal. In the east the 
range does not include Assam, Bhutan, and Burma. Southward it reaches 
the Nilgiri hills or even Cape Comorin [Kanyakumari]. Hyaenas do not 
occur in Ceylon [Sri Lanka]. 

In Africa the striped hyaena was found, except deep in the central parts 
of the Sahara and Libyan deserts, in the northern part of the mainland, and 
south of the Sahara commencing from Azbine (around 19° N. Lat.), in the 
Sudan, Ethiopia (Abyssinia), Somali, Uganda, Kenya, and northern 
Tanganyika. At present, the hyaena is already extirpated in some of these 
areas (Egypt) (Fig. 2). (V.H.) 

Geographic Variation 

Geographic variation of the hyaena is insignificant in spite of the fact that 
a fairly large number of forms have been described (about 24 names). In 
any case the species and subspecies described outside the Soviet Union, 
India, and Iran are not justified; only one form exists throughout the whole 
of India and much, if not all, of the Near East. Its differences from other 
forms, mainly congruent races, have not been satisfactorily explained and 
it is tentatively considered the nominal form. In the Soviet Union, therefore, 
only one subspecies is recognized. 

Asiatic striped hyaena, H. h. hyaena Linnaeus, 1758 (syn. striata, indica, 
zarudnyi, bokharensis, bilkiewieczi, and satunini). 

The description given above pertains to this form. 

This subspecies is found in the Trans-Caucasus (now almost extinct), 
southwest and southem Turkmenia, and plains and slightly elevated regions 
on the right bank of the Pyandzh and upper Amu-Darya from the Darvaz 
mountains to Kugitangtau. 

Outside the Soviet Union this subspecies lives in the Asian part of the 


range, except the southern part of the Arabian peninsula (form syriaca 
apparently does not differ from the nominal form). 

The following forms are usually recognized outside the Soviet Union: 1) 
H. h. syriaca Matschie, 1900 — Syria; 2) H. h. sultana Pocock, 
1934 — southeastem part of the Arabian peninsula; 3) H. h. vulgaris 
Desmarest, 1820 — Egypt and North Africa to eastem Algeria; 4) H. h. 
barbara Blainville, 1844 — western Algeria and Morocco; and 5) H. h. 
dubbah Meyer, 1791 — Sudan, Somali, Kenya, and northern Tanganyika. 
The geographic variation of the species evidently requires revision. 


Population. In the western part of the range within the Soviet Union at 
the end of the last century, hyaenas were extremely rare on the southwest 
coast of the Caspian Sea and nearby mountains, extending north to Derbent 
and Deshlagar (Poletika, 1911; Dinnik, 1914; Heptner and Formozov, 1941). 
31 In the eastem and southern Trans-Caucasus, this carnivore was often 

encountered, but ver}' sporadically. In the 1860s it was "common" along 
the Kur' valley and the adjoining low mountains westward to Tiflis (Tbihsi). 
In those years hyaenas were caught quite frequently around Tiflis, Yerevan, 
and Nakhichevan (Radde, 1899). In the Makhats mountains near Tbilisi 
hyaenas were considered to be "extremely common" in the 1880s. They 
were also common at that time in Karayaz. Early in this century the 
population in this region decreased drastically, and hyaenas disappeared, 
for example from the vicinity of Tbilisi, but were still "quite common" 
in the Shirak steppe and Geok-Chai, Echmiadzin, and Surmalin districts. 
Stray animals were encountered in Zangezur, around Ordubat, near Sal'yan, 
in the lower course of the Kur', and in the Talyshinsk mountains along 
the boundary with Iran (Dinnik, 1914; Satunin, 1915). 

Commencing in the 1930s the hyaena population in the Trans-Caucasus 
continued to fall rapidly and the animal disappeared in many regions. 
Individual animals survived only in the Shchekin and Kartalin plateaus in 
the Shirak and Adzhinaur steppes (Vereshchagin, 1947, 1959), and in several 
other regions in the eastem parts of Georgia and Azerbaidzhan. In 1959 
hyaenas were caught in Azerbaidzhan in the Burun-Guvakh mountains in 
Safaraliev region, where they had never been seen before (Episkoposyan 
and Babakevkhyan, 1959). In the Armenian Soviet Socialist Republic in 
the first half of the twentieth century, the hyaena was encountered very 


rarely. In 1925 it was observed in the neighborhood of Navruzl (Artashat 
region). Later, a single animal was caught in Shamshadinsk region (Dal', 

By 1970 in the Traas-Caucasiis the striped hyaena was still encountered 
singly in the Adzhinaur steppes, in the Kartalin and lori plateaus and also 
in the Araks valley in the Nakhichevan Autonomous Soviet Socialist 
Republic. Sightings and captures were considered a great rarity at this time 
in Gobustan around Shemakha and in Karabakh (Agdamsk, Agdzhabedinsk, 
Zhdanovsk, Fazulinsk, and Dzhabrail'sk regions) (Aliev, 1971). Instances 
are known of their appearance at other places. In 1962 a hyaena was met 
with in the foothills near Geok-Chai; in 1964 in Nakhichevan Autonomous 
Soviet Socialist Republic between Khanagya and Kazanchi villages; in 1967 
in Adzhinaur steppes, in the Nukhinsk region, and the environs of Kichik- 
Dakhna village; and in the summer of 1969 in Talysh, Zuvanda, and near 
the village of Gosmal'yan, Leriksk region. Solitary animals are still sighted 
in these places from time to time (Kh.M. Alekperov). Even in the past the 
extreme rarity of the hyaena was noted in Talysh and Zuvanda (Burchak- 
Abramovich, Mamedov, 1966). 

The total population of the hyaena in the Trans -Caucasus in 1968 was 
placed at less than 150 to 200 animals (Aliev, 1971). In eastern Georgia 
they are "on the verge of extinction" (Arabuli, 1970). In the Trans-CaucasiLS 
26 hyaenas were caught from 1930 to 1940, and not more than 5 or 6 from 
1940 to 1950 (Vereshchagin, 1959). From 1951 through 1958 on the territory 
of Azerbaidzhan only four hyaenas were caught, of which two were from 
Yevlakhsk region, one from Agdzhabedinsk, and one from Gednidsk (Aliev, 
1971). In the last 12 years no skins of this predator have been received 
in the processing units of Azerbaidzhan, although some animals were caught 
during those years (Ю1.М. Alekperov). 

In southwest Turkmenia small numbers of hyaenas used to be sighted 
on occasion in the Atrek valley, and less often along the Caspian Sea coast. 
For example, near FCara Degish and Chalayuk the hyaena has been reported 
only a few times in the region of Lake Bol'shoi Delil and Kara-Bab and 
Kayalydzh mounds. Sightings occurred regularly in these places until 1913, 
were again reported between 1917 and 1922, and then not until 1941. Near 
Gasan-Kuli two hyaenas were caught in 1938 and one in 1941. One or two 
animals were caught at the same place in subsequent years also. The hyaena 
was seen slightly more often on the Messeriansk plateau (north of the lower 
Atrek; Dement 'ev, 1948, 1955; Samorodov, 1953). 

In the Kopet-Dag and its foothills the hyaena was not rare but was 
encountered everywhere, going out also on the adjacent plains. Reports of 
some nineteenth century researchers (Radde and Wal'ter, 1889; Zarudnyi, 
1890; Varentsov, 1894) about the rarity of hyaenas in this region hardly 


reflect the actual situation or indicate a temporary phenomenon. In the first 
two decades of this centur}' the animal was commonly encountered around 
32 Ashkhabad itself, in Gindovar gorge, and in Bagir village (information given 
to V.G. Heptner by S.I. BiFkevich). In the 1920s, 1930s, and even 1940s 
it was common enough, though numbers were not great; in all of the Kopet- 
Dag mountains and, perhaps, from this area, the predator found its way 
into Uzboi. At the end of the 1920s a hyaena lair was excavated in the 
foothills east of Ashkhabad, almost on the plains (V.G. Heptner). 

The hyaena was rare in much of the territory of the Murgab basin, 
especially in the northern part (Flerov, 1932). At the end of the 1930s and 
in the 1940s it was, however, quite common on the upper Murgab and along 
the Kushka, and some skins (about 10 in 1942) were received in Sary- 
Yazy for curing every year (V.G. Heptner). In the 1950s the animal became 
relatively common even in the north. For example, a hunter from the 
Bairam-Aliisk region caught 18 hyaenas in a few years and 4 in the winter 
of 1957-1958 (raeshchinov, 1958). 

In the extreme southeast of Turkmenia, at Badkhyz, the hyaena was 
"a quite common animal" until recently. It was often sighted even in regioas 
adjoining Badkhyz on the north, northeast (Karabil' upland), and east. Several 
were caught every year in the environs of Kala-i-mor on the Kushka River. 
The hyaena was likewise common in the mountains of Gyaz'-Gyadyk at 
Tedzhen and along the Tedzhen at Serakhs as well as above and below 
it (Heptner, 1956). 

The hyaena had become quite rare by 1962 in the low hills of Badkhyz 
and adjoining regions: during daily excursions for 47 days the predator was 
sighted only once. Six of the dens discovered were uninhabited (A.A. 
Sludskii). In the low mountains of Gyaz'-Gyadyk, however, this animal 
was not particularly rare (Yu.K. Gorelov). 

By 1970 the hyaena had become rare even in the eastern part of the 
Karakum close to Karakum canal, where it had persisted at several places 
in the 1940s and 1950s [Zenkush (Lengych), Neder-Belent, Kert-Kuyu, 
Aitysh-Kuyu, and others], in fact, it was "a commonly occurring" large 
predator (Sekunova et al., 1956; Nur-Gel'dyev, I960). 

Over the last 12 years (to 1971) in Turkmenia, only one hyaena skin 
has been cured although many animals were caught; skins were not sent 
for curing because they fetched a very low price. On the whole, in Turkmenia 
during the last 40 to 50 years, the hyaena population and its fluctuations 
depended on the population of ungulates, primarily arkhar sheep and goitered 
gazelle, and wolves, who killed these ungulates. The disappearance of 
goitered gazelle and reduction in the population of mountain sheep in the 
last decade have determined the fate of the hyaena (V.G. Heptner). 

The hyaena was rare even in the 1930s to the 1950s in the valleys, 


foothills, and low mountains of the ranges of southwest Tadzhikistan (Rerov, 
1935; Cherayshev, 1958). In the 1960s the population in this region 
decreased even more. During nocturnal observations on predatory animals 
at water holes and their habitats for over three years, the hyaena was 
encountered only twice in the Karshitau (Koktau) range at Koktash in the 
Leninsk region (Arutyunov, 1964). A hyaena was sighted in September, 
1969, on the northern slope of this same range (Sambuli), and on its southern 
slope (Lyaur) in September, 1970 (A.K. Kovalev). In 1967, a young female 
was caught in the Babatag range (Isambai) (A.I. Kirkhadzhi). Hyaenas were 
sighted a few times in the Garua-Niushti range in September, 1965, 1966, 
and 1967 (Dagana-Kiik, Isambai; A.K. Kovalev). In Tadzhikistan not more 
than 5 to 12 hyaena skins were cured annually from 1932 to 1949. Between 
1949 and 1960 three hyaenas were caught alive (D.P. Dement'ev). From 
1953 through 1963 only five skins were cured; the animals had been caught 
in Ordzhonikidzeabad (1955), Kuibyshev and Dangarinsk (1960), and the 
Pyandzhinsk regions (1963). No skins were received for curing from 1964 
through 1970. ^ 

In Uzbekistan the hyaena was rare in the lower course of the Shirabad 
River, in Shirabadsk region, although it probably occurred in the adjoining 
Termezsk region in "large numbers on Amu-Darya shore" (Sultanov, 1939); 
the latter is extremely dubious. The hyaena was rare along the Surkhan- 
Darya and in the adjoining mountains. Farther north and eastward, it was 
absent. By the 1950s the hyaena had become "a very rare, almost extinct" 
animal in Uzbekistan (Ishunin and Lustin, 1959). 
33 Evidently the hyaena is fewer in numbers today in most places in 

southern Turkmenia and in regions adjoining Afghanistan and Iran. In the 
Trans-Caucasus, southwest Tadzhikistan, and southeast Uzbekistan, the 
hyaena is rare and its population has rapidly declined over the years. Soon 
it will be extinct, especially in the Trans-Caucasus and Uzbekistan. 

Outside the Soviet Union the hyaena is quite rare in the northern parts 
of Iran and Afghanistan. It is still common in Baluchistan and numerous 
at some places in Hindustan (Pocock, 1941; Prater, 1965; Lay, 1967). 

In Africa the striped hyaena is now rare in northern areas, i.e., Egypt, 
Libya, Tunisia, and Morocco. It is absent altogether in the Sahara but more 
common in the savannas of central Africa; it is quite common at some places 
in eastern parts of Africa (Sudan, Somali, Kenya, and others). In central 
and east Africa the striped hyaena inhabits the same regions as the spotted 
hyaena (Crocuta crocuta), but at places where the latter is common, the 
former is rare, and vice versa (Ronnefold, 1969). 

Habitat. In the Trans -Caucasus the hyaena inhabits sections of hilly 
plateaus (lori and KartaUnsk) frequently occupied by mountain steppes, clay 
ephemeral deserts or semideserts, here called "steppes," for example Shirak, 



Mugansk, Adzhinaur, and others, and is confined particularly to the 
surrounding rugged foothills (bozdagi) with dry stream beds, erosion 
channels, and ravines. It also inhabits low, eroded clay mountains with desert 
or steppe vegetation, overgrown in some sections by juniper and pistachio; 
for example, the desert region of the Bozdag range. At places it ascends 
the mountains to heights of 1,800 to 2,000 and 2,100 m above msl (Kyal'vyaz 
in the Talyshin mountains; Dalidag near Istis, the Kel'badzharsk region 
of the Azerbaidzhan Soviet Socialist Republic; F.F. Aliev). In general, 
however, the hyaena avoids high mountains. It also does not live in dense 
forests, but is encountered in small groves and floodplain thickets on rivers, 
for example the Araks and Kur', but rarely in gardens and vineyards. It 
has often been caught on the sea coast, for example at Vzmorsk station 
near Deshlagar and Derbent (Dinnik, 1914; Satunin, 1915; Aliev, 1971; 
Kh.M. Alekperov). 

In Turkmenia the hyaena lives along the stony desert foothills intersected 
by narrow valleys with sparse woody vegetation, arroyos, scarps, ап(^ along 

33 Fig. 7. Gorge in the Gyaz'-Gyadyic mountains (Turangovoe in the Kerlek area), 

habitat of hyaena and arkhar sheep (Oxas amnion cycloceros). Leopard and wolf 
also live here. Pistachio trees {Pistacia vera) grow along the slopes of the gorge. 
Badkhyz preserve, southern Turkmenia. May, 1962. Photograph by A.A. Sludskii. 


rocky gorges almost devoid not only of woody and shrubby vegetation, 
but also grassy vegetation (Kopet-Dag, Gyaz'-Gyadyk, Badkhyz, Karabil') 
(V.G. Heptner, 1956). It likewise occurs deep inside the mountains 
throughout the Kopet-Dag, but rarely and only at some places in the sandy 
desert of the Karakum, and more often in riparian thickets along rivers, 
for example among tamarisk at Kushka and tamarisk and poplar tugais at 
Tedzhen (V.G. Heptner). 

At Badkhyz, in southeast Turkmenia (about 850 m above msl), the 
hyaena inhabits the Er-oilan-duz and Namak-Saar basins and also Kyzyl- 
Dzhar plateau in the rolling countryside where high ridges alternate with 
broad open valleys, and very narrow and deep erosion channels. At places 
in these lowlands, weakly saline to saline springs are seen. Along the tops 
of the high ridges, and on their northern slopes, grow large, solitary trees 
of pistachio with crowns in the form of a dark green tent. Over the years 
the sandy loam soil of the ridges and valleys has become densely overgrown 
with a tall herbaceous caфet of meadow grass {Poa bulbosa), desert sedge 
{Carex pachystylis), and different varieties of sagebrush [Artemesia]. 
Gigantic fennels (Ferula badrakema) with peduncles as thick as a hand 
and reaching a height of up to 2.0 m are typical. In places, during years 
of good rainfall, they grow so densely that they impede the movement of 
man, and their thickets create the impression of a "forest". In some sections, 
above the low herbaceous cover, thickets of other large umbellifers grow, 
such as dorema (Dorema aitchisonii), the much smaller cousinia, bushes 
oiCoIligomim setosum, and black saxaul. This type of landscape is usually 
called a semisavanna.^ 

Among vertebrates, reptiles of various species are found in large 
numbers in Badkhyz, in particular monitors and steppe tortoises. In some 
years both large and red-tailed gerbils are abundant, while thin-toed ground 
squirrels and sand hares are encountered occasionally. Even in the 1940s 
there were innumerable goitered gazelle, common mountain sheep [sic], 
and kulan. Numerous flocks of bCarakul sheep are seen throughout the year. 
35 Hyaenas hunt some of these animals or feed on their carcasses. Large 
predators capable of competing^with hyaenas, i.e., wolf, cheetah, and leopard, 
are rarely seen. On Kyzyl-Dzhar plateau in May, 1962 hyaenas were sighted 
only once in 20 days of daily trekking, but the animal's characteristic feces 
were detected several times. 

In Badkhyz, however, the hyaena also inhabits the low mountains of 
Gyaz'-Gyadyk, which are intersected by deep valleys of small rivers with 
outcrops of limestone and sandstone. The steep slopes of these valleys are 

-'For photographs depicting the nature of the Badkhyz environment, also see sections 
on cheetah, leopard, and other species. 


34 Fig. 8. Rocky trail along bottom of a gorge used by hyaenas and also by porcupine, 

arkliar sheep, and other animals, sometimes even leopard. Kerlek region in Gyaz'- 
Gyadyk mountains, Badkhyz preserve, southern Turkmenia. February, 1965. 
Photograph by V.G. Heptner. 

covered with desert sedge and herbs. Along the gorges (down to the very 
bottom) of northern slopes grow centuries-old pistachio trees with trunks 
up to 1.0 m in diameter but a height of only 5.0 to 6.0 m; along the bottom 
of the gorges wild fig trees abound. Small saline springs occur in some 
gorges. In the larger gorges, for example in Kerlek, a narrow strip of reeds 
grow along the saline river bed. Hyaena tracks have been found at some 
of these springs as the animals use them as water holes and set up their 
dens in nearby rock fissures. 

The striped hyaena is also encountered in the deep basin (to 500 m) 
of Er-oilan-duz with a steep northern arroyo, and an undulating bottom 
relief. Low mud cones of greatly disintegrated volcanic rocks rise from it. 
The central portion of the basin is covered by extensive solonchaks. Along 
the edges of the basin grow small thickets of white saxaul, saltwort, and 
other plants. The grassy cover consists of desert sedge, sagebrush, and 
halophytes. During a week-long excursion, two fresh tracks of hyaenas were 

28 ■ 

detected in this basin and two old burrows and a den found under a projecting 
rock among mud volcanoes.'^ 

In southwestern part of the Karabil' upland, the hyaena inhabits 
extremely rugged, almost inaccessible, and unpopulated sites (Sheram- 
Kuyu, ShikhmuUa, and Darvaza-kem wells, and many others), where flocks 
of Karakul' sheep are driven for pasturing in autumn and winter (Nur- 
Gel'dyev, 1960). 
36 In Turkmenia, however, the hyaena inhabits the extensive sandy desert, 

for example in the southeastern parts of the Karakum, where it is common 
in the black saxaul thickets at Lengych, Neder-Belent, Kert-Kuyu, Aitysh- 
Kuyu, and other wells at which it regularly or in some seasons snatches 
sheep (Sekunova et al., 1956; Nur-Gel'dyev, 1960). Thus on January 30, 
1956, in the sands around the Lengych well in a trough 3.0 km x 3.0 km 
densely overgrown with black saxaul to a height of 3.5 to 4.0 m and thickness 
30 to 40 cm, a pregnant female hyaena was caught. In this area hyaenas 
once lived rather permanently and were regularly seen from 1948 (Sekunova 
et al., 1956). 

35 Fig. 9. Lake Er-oilan-duz basin, habitat of hyaena. Arkhar sheep, kulan, and goitered 

gazelle are also found here. Badkhyz preserve, southern Turkmenia. May 1962. 
Photograph by Yu.K. Gorelov. 

''Data for Badkhyz are from V.G. Heptner, 1956 and unpublished material 
of A.A. Sludksii (1962) and V.G. Heptner (1942, 1948, and 1%5). 


In the extreme south of Uzbekistan and in southwest Tadzhikistan, the 
hyaena was confined roughly to the same biotopes as in Turkmenia. 
Moreover, in this region, h was noticed regularly in tugai thickets consisting 
of turanga, oleaster, tamarisk, giant plume grass, and reeds. Thus at the 
southern extremity of the Kugitangtau range, on the low mountain of 
Kulanashar, watered by the Amu-Darya, the hyaena was confined to tamarisk 
shrubs. Farther north this animal was found in the mountains at Kyzylalma 
village, in montane steppe with innumerable dryland and irrigated farms 
(Zarudnyi, 1917). 

In the southern spurs of the Gissar range, the hyaena inhabits the foothills 
of low mountains (Babatag, Aktau, Karshitau [Koktau], Garda-Niushti, and 
others) with steep forest -covered slopes and extremely rugged gorges and 
ravines, at the bottom of which saline springs are sometimes visible. The 
northern slopes are covered with steppe vegetation, and the southern with 
semidesert. There is a pistachio plantation in Babatag. The vertebrate fauna 
of these mountains is poor (steppe tortoise, chukar and desert partridge, 
porcupine, mountain sheep), but [domestic] sheep are abundant (Arutyunov, 
1964; A.A., Sludskii; A.K. Kovalev; A.I. Kirkhadzhi). 

In the tiger sanctuary ' 'Tigrovaya Balka' ' on the lower reaches of the 
Vakhsh, hyaenas regularly inhabit floodplains and different types of tugai 
vegetation. The highest elevations of the floodplains, sometimes with sand 
dunes, are overgrown with turanga trees. The trees in such a forest are sparse, 
seedlings and undergrowth are almost absent, and the herbaceous cover 
poor. In a turanga tugai one meets with small depressions and dry river 
beds, usually densely overgrown with tamarisk and some individual oleaster 
trees. Extensive depressions, periodically inundated by floodwaters, are 
densely covered with giant plume grass and more rarely by huge tamarisk 
bushes, which form almost impenetrable jungles. In the Vakhsh floodplain 
are many large fresh-water meanders and streams, the banks of which are 
densely overgrown with tugai or massive reeds and cattails. From the west, 
spurs of the Akatau range run almost up to the floodplain, and from the 
east, the Kashkakum sand desert. The vertebrate fauna in ' Tigrovaya Balka" 
sanctuary is very rich (monitor lizard, steppe tortoise, black-breasted 
pheasant, many ducks, sand hare, porcupine, coypu, goitered gazelle, and 
tugai deer). Many cattle and horses are here throughout the year. In the 
adjoining desert are steppe tortoise, gerbil, thin-toed ground squirrel, and 
other animals, likewise encountered in the floodplains, abound. In the sand 
along the floodplains [domestic] sheep are numerous (A.A. Sludksii). Two 
or three pairs of hyaenas regularly inhabit "Tigrovaya Balka" (46,000 
hectares) (A.I. Kirkhadzhi). 

Near the Pyandzh River hyaenas were encountered in a narrow valley 
between the end of the infertile Takakamar range and the river. Dense 


tamarisk shrubs cover this gravelly soil impregnated with salt and at places 
filled with sand or broken rock (Zarudnyi, 1917). In Amu-Darya valley, 
in the region of Termez, this predator was formerly rather often encountered 
in reed beds and in ravines in adjacent river bank terraces (Suhanov, 1939). 

Thus throughout the Soviet Union the hyaena mainly inhabits foot- 
hills, clay ephemera deserts of the southern type, and montane steppes 
characterized by rugged topography and an abundance of dry river beds, 
gorges, precipitous scaфs, and caves to serve as shelter. More rarely, the 
hyaena lives in sandy deserts, tugais, etc. The animal avoids high mountains 
and dense forests. Habitat selection largely depends on abundance and 
37 accessibility of food — carcasses, fresh bones of wild and domestic ungulates, 
live ungulates, and steppe tortoises, and other animals. 

The hyaena, with its poor coat of sparse guard hair, thin underfur, and 
large ears almost devoid of hair, is little adapted to low atmospheric 
temperatures and cannot survive cold below -15 to -20°C. Hence it is 
distributed south of the January isotherm of 1.0°, does not extend high into 
the mountains, and inhabits only regions with a frostfree period of 230 to 
270 days. 

The plains and low mountain regions of the Trans-Caucasus, Atreksk 
basin, the Kopet-Dag foothills, Tedzhen valley, the Murgab, upper part of the 
Amu-Darya valley and lower Pyandzh, Vakhsh, and several other rivers are 
prominent in the Soviet Union for length of frostfree period above freezing 
(Freikin, 1954; and others). The range of the hyaena within the Soviet Union 
is confined to these territories, and the animal seldom ventures out of them. 
Only on the west banks of the Caspian Sea, where the climate is mild because 
of the proximity of that water body, does the hyaena penetrate somewhat 
farther north. The absence of hyaenas in the northern half of the Karakum 
and the central and lower courses of the Amu-Darya is explained, apparently 
by prolonged freezes and short fi-ostfiree periods. To the north of the Karakum 
frosts of up to -20°C occur even in November, the temperature dropping 
sometimes to -25°C in January, and exceptionally even as low as -33°C. 
The mean January temperature is -2.0°C. The coldest weather occurs in 
the delta of the Amu-Darya where, at Nukus, the mean January temperature 
is -5.4°C. Within the range of the hyaena, at Termez the mean January 
temperature is 1.6°C and in extreme southern Turkmenia, even 4.7°C. The 
absence of permanent snow cover is also typical of hyaena habitats. 

In India, Pakistan, Afghanistan, Iran and Africa, the hyaena inhabits 
open desert expanses, especially in stony, rugged ravines. In Africa the 
animal is common even in the savannas. In forests it is restricted to edges 
or remains close to roads and paths (Pocock, 1941; Bourliere, 1955a; Prater, 
1965). In Iran its vertical distribution extends up to 2,250 to 2,475 m above 
msl (Dinnik, 1914; Lay, 1967). 


Food. Precise information on the food of the hyaena within the Soviet 
Union is scant. It is regarded everywhere as a carrion feeder, consuming 
mainly the carcasses of ungulates in different stages of decomposition, fresh 
bones, cartilage, ligaments, and bone marrow. The hyaena cmshes long 
bones into fine particles and swallows them. Sometimes, though, whole 
bones or large pieces are found in its stomach. In the Trans-Caucasus, apart 
from carrion, it steals dogs, sheep, and other small domestic animals. Whether 
it attacks wild animals and birds in this region is not known (Dinnik, 1914; 
Satunin, 1915). In summer it consumes locusts and reptiles (Dal', 1954). 

In a hyaena burrow in Badkhyz five horns of goitered gazelle were 
found, as well as a lower jaw and long bones, together with the shells of 
small steppe tortoises and the paws of red fox. Bones of a young wild boar 
were scattered in another four lairs and bones of mountain sheep in a fifth. 
In yet another hole the shriveled leg of a young kulan (onager) was found, 
and in still another the old bones of a mountain sheep were scattered. It 
is possible that the hyaenas did not catch these ungulates themselves, but 
merely picked up their bones (A.A. Sludskii). In one den the skull of a 
mountain sheep, bones of horses, and other bones were discovered. The 
stomach contents of one of two hyaenas caught contained the remains of 
a calf (evidently carrion), while the other contained a large porcupine quill 
broken in two (Yu.K. Gorelov). The stomach of a hyaena caught on January 
30, 1956, in the southeastern part of the Karakum contained a pair of hooves, 
bones, and wool of a domestic sheep, and bits from the shell of a steppe 
tortoise. A male (in southwest Karakum) (of a pair, of which the female 
was caught) quickly bit to death two Karakul sheep (Sekunova et al., 1956). 
Similar information is available to show that in Turkmenia the hyaena feeds 
not only on carrion, but also hunts dogs, monitor and other lizards, and 
rodents, and coasumes steppe tortoises, insects, and musk- and watermelons. 
In a single visit to a melon patch, this animal has been seen lopping off 
six or seven large watermelons (Heptner, 1956; Shcherbina, 1970). 
38 In southwest Tadzhikistan (Karshitau range), a young male caught on 
September 3, 1963 yielded stomach contents weighing 600 g, consisting 
of three large steppe tortoises, a small bit of skin with domestic goat wool, 
ten locusts, caper fmits, and a small amount of leaves and stalks (Anityunov, 
1964). The stomach of a young female caught on August 18, 1967 in the 
Babatag range contained the remains of 3 or 4 steppe tortoises, 18 locusts, 
and dry grain; the weight of this stomach was 300 g (A.I. Kirkhadzhi). 
In the tugai along the Pyandzh, fecal exarninatioas revealed drupes of oleaster 
{Elaeagnus angiisti folia). The number of hyaenas increased in the period 
of ripening of these fruits in October-November in the tugais. These fruits 
are also consumed in large numbers by jackals, foxes, and jungle cats 
(Chernyshev, 1948). 


In Iraq instances are known of this predator attacking horses and asses 
and also feeding on tortoises (Khett, 1959). 

In India the hyaena feeds mainly on carcasses of animals dead from 
diseases or killed by other predators; it also attacks dogs, sheep, goats, and 
calves (Jerdon,"' 1874; Pocock, 1941; Prater, 1965). Many instances of 
hyaenas chasing leopards away from their quarry have been recorded 
(Pocock, 1941; Prater, 1965). 

In Africa carrion constitute the main food of the hyaena. It also feeds 
on large lizards, snakes, and fruits. It attacks, but not often, sheep, goats, 
and dogs (Ronnefeld, 1969). 

It is possible that the striped hyaena attacks live animals far more 
frequently than supposed. The animal's nocturnal mode of life and innate 
caution make it difficult to assess the degree to which it is predatory, as 
is known in particular for the wolf in the Soviet Union. For example, it 
was long thought that the spotted hyaena (Crocuta crocuta) was a typical 
scavenger, but recent observations have established that it feeds mainly 
on carrion, only when living close to inhabited villages or in places where 
large predators are numeroiLS. Elsewhere it actively and successfully attacks 
adult zebra, gnu, gazelle, or their young. In some regions of Africa hyaenas 
have destroyed up to 50% of the newborn antelope calves and older juveniles 
(Kruuk, 1966). 

Frequent finds of shells of steppe tortoises in hyaena stomachs in 
August -September and even in January when they are in summer- winter 
hibernation, suggest that the predators somehow detect the tortoise's 
hibernacula and dig them out. Throughout most of the range of the striped 
hyaena, steppe tortoises are extremely abundant and constitute a ready prey 
for the carnivore. In southwest Tadzhikistan, after tortoises had gone into 
hibernation, one hyaena succeeded in capturing three after 2.5 hrs of 
nocturnal hunting around a water hole. Obviously, tortoises are an important 
coastituent of the diet of the striped hyaena, especially when other sources 
of food are scarce. 

In zoological gardens the hyaena receives a daily ration of 3.0 kg of 
meat with bones (Balaev, 1940). 

Feeding mainly on carrion and bones, the hyaena evidently requires 
more water to quench its tliirst than other carnivores. In southeast Turkmenia 
hyaenas drink from rivulets, springs, and wells in which water is close to 
the .soil surface. In Badkhyz (southern Turkmenia) they are closely confined 
to saline springs; a liter of this water can contain up to 20 g of salts. The 
animals drink this water in the warm period of the year (A.A. Sludskii). 
Hyaenas inhabiting the southea.stern parts of the Karakum are usually 

*Not in Literature Cited — Sci. Ed. 


confined close to wells and drink water remaining in troughs after sheep have 
dnink (Sekunova et al., 1956). In southwest Tadzhikistan this carnivore drinks 
from rivulets, canals, and irrigation ditches; at places where fresh water 
is not available, it drinks from springs of saline water. It usually remains 
close to such springs in the warm season of the year. In winter and spring 
the animal drinks irregularly and, during this period, may be encountered 
15 to 20 km away from the nearest water hole (Yu.K. Gorelov). Evidently, 
however, it will migrate long distances in search of water (Flerov, 1932). 
39 In the Kushka region of Turkmenia during the ripening of water- and 
muskmelons hyaenas regularly devour them, but they prefer the first, for 
which they enter the melon patch. Watermelon is mainly consumed for the 
риф08е of quenching thirst (Heptner, 1956; Shcherbina, 1970). Water- and 
muskmelons were avidly consumed by a hyaena held in the Tashkent 
Zoological Garden (Balaev, 1940). 

Home range. The size of the breeding area during the reproductive period 
is not known. In search of prey a hyaena may wander far from its den. In 
"Tigrovaya Balka" sanctuary in Tadzhikistan two to three pairs of hyaenas 
ranged over an area of 46,000 hectares. Outside of the breeding period, 
hyaenas are probably nomadic. However, in Turkmenia instances are known 
of great attachment by the hyaena to a given site. Thus, one hyaena lived 
for several years in a single lateral fissure of Gindovar gorge in the Kopet- 
Dag in the vicinity of Ashkhabad in a relatively populated locality. It is 
quite likely that such instances are due to the availability of convenient 
and safe hiding places, which by and large are few. The animal may be 
confined to a given place even after the mating season (V.G. Heptner). 

The hyaena marks its territory by leaving feces at dens, burrows, and 
in elevated places, such as the summits of hills and ridges along trails. 
Because of their high calcium content, the feces turn white rapidly and are 
visible from fairly long distances. Moreover, secretions from the special 
preanal scent glands of the hyaena also impart to the feces a characteristic 
odor. Such feces thus serve as scented and colored marks. 

Burrows and. shelters. Throughout its home range, the hyaena establishes 
lairs in caves, rock fissures, and erosion channels, or in burrows formerly 
occupied by the porcupine or wolf, and also digs its own dens. Whatever 
previous animal may have tenanted an erstwhile hole, when a hyaena takes 
over occupancy, a characteristic collection of bones of wild and domestic 
animals will be found at the entrance (V.G. Heptner; Fig. 11). A burrow 
uncovered at Kumkulab (not far from Gurmab) in the Kopet-Dag was located 
in the central part of the broad valley of the Mergen'ul' River. The entrance 
to the small cave, about 2.5 m high, was under a vertical wall of low rock. 
Farther in the cave narrowed and ran deep inside in the form of a tunnel. 
The lair was not visible from outside. At the entrance to the burrow and 


among the rocks, many large bones of several species of animals were 
scattered in a semicircle 25 m in diameter; most were old and sun-bleached. 
They stood out sharply against the background of stones and the location 
of the burrow was visible some distance away. It was not visible from the 
road, however. The bones were mostly those of domestic animals (horses, 
cows and asses) but a few bones of mountain sheep were also present. The 
burrow had been used for several years (V.G. Heptner). In loose soil in 
ravines of the Kopet-Dag foothills, the hyaena sometimes digs its own 
burrow. The entrance is usually broad. In Turkmenia hyaenas often use 
the burrows of porcupines (V.G. Heptner). 

Six of the lairs examined in Badkhyz (southern Turkmenia) were 
identical to the one described above. One burrow was located on a completely 
exposed crest of a gentle ridge overgrown with low grass and solitary bushes. 
The hole had two entrances 60 cm x 40 cm. At both entrances there were 
large soil heaps measuring several cubic meters. Around the hole there were 
many feces in specially dug small pits. Scattered nearby were five pairs 
of horns of goitered gazelle, their long bones, tortoise shells, fox paws, 
and the skull of a porcupine. Another four lairs were found in broad fissures 
of limestone outcrops in a fairly deep gorge in the bed of the saline Kerlak 
River, on the northern slope of which grew old pistachio trees and at the 
bottom wild figs. At the entrance to one shelter lay the bones of mountain 
sheep and a juvenile wild boar. A lair found in the Er-oilan-duz basin was 
located on top of a small conical hill in a broad fissure among the outcrops 
of volcanic rocks. In addition to the main entrance this burrow had two 
exits among the rocks; around these lay the old bones of a mountain sheep 
and the legs of a young kulan. A few hundred meters from this lair, in a 
niche under a rock, was another hyaena den containing wool and bones 
of mountain sheep (A.A. Sludskii). At the same elevation hyaenas often 
occupied the holes of porcupines after widening them (Yu.K. Gorelov). 
42 In the southeast Karakum a hyaena burrow was found at the edge of 
a dense thicket of black saxaul. The entrance lay under a projection of 
compressed sand at the rim of an old well and was 67 cm wide. The hole 
ran to a depth of 3.0 m and extended in length for 4 m 15 cm. There were 
no chambers or enlargements inside. A second hole (2.0 km from Chaerla 
well) lay directly on the path leading to the well. The entrance to this was 
protected by low saltwort and saxaul thickets. Hyaena feces were evident 
around the hole and there were hairs at the entrance. The dimensions of 
the entrance were 72 cm x 38 cm. This burrow extended over 5.0 ra in 
length and 2.5 m in depth (Sekunova et al., 1956). From time to time, in 
dense growth of bushes or reeds, the den for cubs will be open. 

In India the favorite sites for hyaena dens are caves among rocks 
or holes dug on slopes of hills or ravines. Most often hyaenas occupy and 


40 Fig. 10. Burrow of a hyaena on a plateau in tlie Kyzyl-Dzhar area. Badkhyz 

preseive. May, 1962. Photograph by A.A. Sludskii. 

enlarge burrows of porcupines (Prater, 1965). In Africa hyaenas sometimes 
occupy burrows of aardvarks and wart hogs (Cotlow, I960*). 

In the Tashkent Zoological Garden, just before parturition, a female 
began to dig a burrow with a male helping her in the task. Hyaenas dig 
the earth with the claws of the front paws and from time to time remove 
the dug-up soil with their hind legs. If small stones were encountered during 
digging, the male would pull them out with his teeth and drag them away 
from the surface. The hole was dug quite rapidly to a depth of about 1.5 
m; the female whelped in it (Balaev, 1940). 

During the day hyaenas hide in caves, niches, and holes, protecting 
themselves from enemies and the sun during summer and low temperatures 
in winter. Although nomadic, the hyaena rests during the day in temporary 
cover, such as ravines, niches, pits, dense thickets or reeds, and plume grass 
or Erianthus. In Badkhyz a hyaena was observed resting during the day 
on an open plateau near a bush (A.A. Sludskii). In the hills around Kushka 

*Not in Literature Cited — Sci. Ed. 


40 Fig. 11. Entrance to burrow of a hyaena on a level site (see Fig. 10). Around the 

bunow can be seen bones and horns of goitered gazelles. May, 1962. Photograph 
by A.A. Sludskii. 

during the day, hyaena sometimes hides under tumbleweed piles transported 
by the wind into hollows (V.G. Heptner). In the Karakura these carnivores 
rest among saxauls, hiding under wind-felled trees (Sekunova et al., 1956). 
In India hyaenas sometimes rest during the day among tall grass, shrubs, 
in sugar cane fields, caves, and under rocks (Pocock, 1941; Prater, 1965). 

Hyaenas often carry their catch to the burrow and consume it at leisure. 

Daily activity and behavior. The hyaena is essentially nocturnal and 
moves out from its den only after the onset of total darkness, usually returning 
to the shelter long before sunrise. It is rarely sighted during daylight hours. 
43 Once in Badkhyz a hyaena approached a hunter waiting in ambush during 
the day; the animal came so close that the hunter killed it with a blow to 
the head with his rifle (Yu.K. Gorelov). 

Hyaenas are cautious but extremely "cowardly." Sometimes, however, 
especially during the night, they exhibit little fear of man. In Kushka instances 
are known when in twilight a hyaena will persistently trail a man along the 
road, keeping only a few tens of steps behind him. Hyaenas (at least in the 



Fig. 12. Rocky gorge overgrown with pistachio, in which was located a hyaena 

breeding burrow. Keriek region. Badkliyz preserve, southern Turkmenia. May, 

1962. Photograph by A.A. Sludskii. 

1940s) often came right up to houses in Morgunovsk village near Kushka, 
howled at their windows, etc. Hungry animals approached carrion very boldly 
(V.G. Heptner). In Badkhyz hyaenas formerly followed border patrols at 
night, within 20 to 50 steps behind them. It is possible that the animals 
were attracted to the viscera and heads of mountain sheep which the soldiers 
discarded from animals sometimes caught incidentally (Yu.K. Gorelov). 
In the event of unexpected danger the hyaena hides; it is very difficult 
to spot the animal in twilight and at night because of its striped coloration. 
The stripes crisscross the entire body surface, and therefore it is rendered 
inconspicuous. "Feigning death" is a characteristic pose of the striped 
hyaena. For example, if its lair is dug up and the hyaena exposed, the animal 
holds its breath, closes its eyes, and pretends to be dead; it can remain in 
that state for several minutes (Bourliere, 1955*). It also goes into this state 
when attacked by dogs from which it is unable to escape; in such a situation, 

*Not in Literature Cited — Sci. Ed. 


41 Fig. 13. Rock (at lower right) under wliich lies the entrance to a hyaena den. Kamennoe 

gorge, Kerlek region. Badkliyz preserve, southern Turkmenia. May, 1962. Photograph 

by Yu.K. Gorelov. 

the hyaena suffers their powerful bites without resistance (Pocock, 1941). 
When the dogs finally leave, it jumps up and runs away. 

A tamed hyaena fondled by its master and sexually aroused everts its 
rectum to a length up to 5.0 cm. Evidently, such behavior is a form of 
communication between these animals since special scent glands producing 
secretions are located at the anal opening. During rut hyaenas evert the 
rectum and press it against prominent objects, thus marking their own 
territory, within which they remain. When an enemy attacks, the hyaena 
similarly everts the rectum and sprays secretions from the rectal glands, 
which give off a strong unpleasant odor (Ronnefeld, 1969). 

While seeking quarry, the hyaena is mainly guided by its olfactory 
sense which, however, is not as well developed as in dogs. Vision and hearing 
are also good although some researchers report that hearing is relatively 
weak (Pocock, 1941). Hyaenas feeding on the remains of prey captured 
by other carnivores, often remain in their vicinity, forming a sort of 
commensal relationship. Such groups of hyaenas are common in places 
in southern Turkmenia where wolves are quite numerous. 


42 Fig. 14. Bones at entrance to a hyaena burrow (see Fig. 15): horses, goitered 

gazelle, arkhar sheep, wild boar, and other animals. May, 1962. Photograph by 

Yu.K. Gorelov. 

There is no information about the methods hyaenas employ for hunting 
live prey in the Soviet Union. In other countries they usually hunt in pairs 
or family groups of five or six animals. They are fleet-footed and hardy. 
Having reached the prey by chasing, the predator grabs it by its flanks, 
in the region of the groin, inflicts large mortal wounds, and pulls out the 
viscera, which causes the victim's death (Krumbiegel, 1954). 

Hyaenas live in pairs, more rarely singly, or form small family groups 
of five or six animals. The pair lives in a single den. 

In captivity hyaenas often quarrel among themselves. One method of 
attack is to attempt to hold the opponent by the leg. In this process the 
attacker approaches sideways. Another method is the throat hold. During 
fights hyaenas often inflict severe wounds on each other, but then upon 
calming dov/n, rest peacefully in one den (Dalaev, 1940). 

Hyaenas caught when young, are easily tamed and behave like dogs 
even after attaining adulthood. They are amenable to training. 

Seasonal migration and transgressions. These aspects are not known. 
In southeast Turkmenia, at Badkhyz, following the drying up of water 
sources, fires, and the migration of wild and domestic ungulates to the 


44 Fig. 15. Rocks among which a hyaena den was found. Er-oilan-duz basin, Badkhyz 

presei-ve, southern Turkmenia. May 1962. Photograph by A.A. Sludskii. 

Kushka valley, hyaenas and wolves follow the migrants. Thus in the second 
44 half of July, 1942 at Morgunovsk village, hyaenas appeared in larger numbers 
than previously. They were sighted very often in tamarisk thickets. These 
carnivores arrived there from the desert. Such a migration of hyaenas to 
this river was not seen in 1941 (Heptner, 1956). The hyaena population 
increased during the autumn in the tugais of the Vakhsh, Pyandzh, and other 
river valleys in southwest Tadzhikistan. This phenomenon may be explained 
by their migration from the surrounding desert. Hyaenas also congregate 
in areas of heavy mortaUty of wild and domestic ungulates during outbreaks 
of epizootic diseases or during heavy sleet. 

In Africa, together with predatory animals such as lions, cheetah, and 
hunting dogs, hyaenas accompany herds of migrating ungulates, attack sick 
and weakened animals, and subsist on the carcasses of the dead. 

Reproduction. The striped hyaena is monogamous. The male helps the 
female in establishing the den and in raising the young. 

The periods of rut reported in literature are only partly true and require 
verification, viz., November- December and January -February for hyaena 
in Trans-Caucasus (Ognev, 1931; Aliev, 1971) and October- November 
in southeast Turkmenia (Shcherbina, 1970). According to observations made 


in the Tashkent Zoological Garden, periods of rut in the striped hyaena 
are not seasonal. A female mated three times in 1937: January 11, April 
15 (three days after the death of her litter), and December 16. She whelped 
on all the three occasions. A female mated twice in 1935: in the middle 
of June and on December 30 (Balaev, 1940). These observations suggest 
that, under natural conditions, hyaena young may be seen in all seasons, 
which is typical of many carnivores in the subtropics and tropics. Coitus 
lasts for over an hour. During mating the male grips the skin of the neck 
of the female in his teeth; it is not restricted to any particular time of the 
day. During the period of mating the animals remain irritable. 

Gestation requires 90 to 91 days (Heck and Hiltzheymer, 1925; Balaev, 
1940; Krumbiegel, 1954). In the Tashkent Zoological Garden a female mated 
on January 22 and whelped on March 16. 

In the Trans-Caucasus cubs appear in April -May (Ognev, 1931). In May 
a pregnant female with four well-developed fetuses was killed there (Satunin, 
1915). In Turkmenia, in the southeastern part of Karakum, on January 30, 
1956, a large pregnant female with considerable subcutaneous and intestinal 
45 fat was killed. She carried three fetuses 9.8 cm in length and 1.0 to 1.5 
months of age (mating had evidently taken place in December; Sekunova 
et al., 1956). In southeast Turkmenia pregnant females were caught in 
January and lactating ones in February (Shcherbina, 1970). In southwest 
Turkmenia hyaena litters were seen in March (G.P. Dement 'ev, 1955). 

The litter invariably consists of one to four cubs, usually three. In the 
Trans-Caucasus tv^'O to four cubs are common (Satunin, 1915); in southwest 
Turkmenia two or three, and in the southeast two to four (Shcherbina, 1970). 
In the Kopet-Dag a litter contained two cubs (V.G. Heptner). In the tugais 
of the Vakhsh valley (southwest Turkmenia), a lair with four cubs aged 
1.0 to 1.5 months was found among reeds in early June, 1950 (Chemyshev, 
1958). In southern Uzbekistan hyaena litters consist of three or four cubs 
(Sultanov, 1939). According to observations in the Tashkent Zoological 
Garden, the hyaena produces one to four cubs, but as a rule three. Thus 
one female in her first whelping birthed a single cub, in the second, four, 
and subsequently three each time (Balaev, 1940). In India hyaena litters 
consist of two to four cubs (Blanford, 1888; Pocock, 1941) but three or 
four in different parts of Africa (Ronnefeld, 1969). 

Whelping lasts, on average, for two hours, commencing usually between 
10:00 a.m. and 12:00 noon. 

Growth, development, and molt. Young are born blind, with closed 
ear passages, and are the size of the pups of an average-sized dog. The 
hair coat of the newborn is whitish-gray with distinct black stripes. Exposed 
to the light, cubs immediately whine like pups and constantly crawl around 
the den. They begin to see clearly on the seventh or eighth day. By the tenth 


day their measurements are as follows: female No. 1 — body length 30 cm 
and tail length 7.0 cm; female No. 2 — 29 and 8.0 cm; and one male 29 
and 8.0 cm respectively. Up to the age of one month the young are confined 
to the den and only later begin to venture out. At first they are very cautious 
and hide in the den at the slightest sound. They emerge from it singly, mainly 
on the leeward side, and before leaving peep out to examine the surrounding 
area. In sheltered localities in the Kopet-Dag, growing cubs often play around 
the burrow during the day (V.G. Heptner). 

The young, up to the age of two months, feed exclusively on their 
mother's milk and begin to take meat only thereafter. The prolonged period 
of lactation is an adaptation since cubs, in their early life, cannot feed on 
carrion, especially bones. 

Both parents participate in feeding the young, by the female more so 
and for a longer period. Before the young emerge from the den the female 
remains with them continuously; if she leaves them at all, it is only for 
a very short while. She is fed during this period by the male. In the presence 
of the female the male will not approach the cubs since she invariably drives 
him away. In her absence, however, the male sneaks to the cubs, sniffs 
them, and licks them. When the cubs begin to emerge from the den, the 
female follows them vigilantly. She does not allow them to remain outside 
for long and, picking them up by their neck with her teeth, pushes them 
back inside the den. In spite of the male's solicitous attitude toward the 
cubs, the female is suspicious of his intentions and invariably chases him 
away whenever he approaches her litter. The female eats the feces of her 
cubs (Balaev, 1940). 

By autumn young hyaenas are roughly one-half the size of their parents. 
Thus a young male caught in southwest Tadzhikistan on September 3, 1956, 
aged about 5 months, had the following measurements: body length 94 cm, 
tail 38 cm, ears 1 1 cm, and weight 19.4 kg. Its deciduous teeth had already 
been replaced by permanent ones and the premolars lost (Arutyunov, 1964). 
A young female caught in the same region on August 18, 1967 had a body 
length of 91 cm, tail 36 cm, ears 11.7 cm, and weight 17.2 kg (A.I. 
Kirkhadzhi). In India an adult male of 150 cm body length weighed 38.5 
kg, and a female 34 kg (Prater, 1965). 

The period of the onset of sexual maturation in the hyaena has not 
been accurately established. In the Tashkent Zoological Garden a female 
bom in 1929 produced her first litter in 1933, i.e., in her fourth year (Balaev, 
1941). Hyaenas live for an average of 12 years but have survived for 23 
in a zoological garden (Rower, 1931). 
46 Molt has not been studied. 

Enemies, competitors, diseases, parasites, mortality, and population 
dynamics. Within the Soviet Union wolves are evidently enemies of the 


hyaena and, until recently, so were tigers and leopards. The main competitors 
for their food are wolves, jackals, red foxes, rarely corsac foxes, and griffon 

The importance of griffon vuhures where they are abundant, for example 
in Badkhyz (in the 1940s), is paramount. Carrion remaining at dawn is 
consumed by vultures in the early hours of the morning, leaving no remnants 
except bones. The following example reveals the competitive relationship 
between hyaenas and large carrion-eating birds, primarily griffons and other 
vultures. In Badkhyz preserve in the early 1940s, when there were large 
numbers of these birds, hyaenas were also common. Goitered gazelle served 
as the major food source for all. After the 1950s and early 1960s goitered 
gazelle became almost extinct in southern Turkmenia and by 1965 nesting 
colonies of griffons had decreased sharply and hyaenas had become almost 
extinct (V.G. Heptner). 

At the same time, as previously mentioned, hyaenas often become the 
commensals of tigers, leopard, and cheetah, since they finish off the remains 
of carrion, especially bones, left behind by these predators. While wolves, 
like hyaenas, crush most bones and consume them, large cats do not. 

Diseases and parasites are almost unstudied. It is known only that 
hyaenas are highly resistant to various infectious diseases and suffer only 
rarely even when held in zoological gardens (Balaev, 1940). A hyaena caught 
on January 30 in the southeast Karakum was infected by Taenia pisifonnis 
in the abdominal cavity; fleas {Synosternum paUidus) were also collected 
from the same animal. The substratum collected from the den of a hyaena 
in the same region at a depth of 3.0 m revealed the presence of mites 
(Eulaelaps stadularis and Haemalaelaps longipes) and ixodid ticks 
{Hyalomma a. asiaticum) (Sekunova et al., 1956). From an animal killed 
in Badkhyz on January 23, 1960, two Ixodes ticks and five Pulex irritans 
fleas were removed (Yu.K. Gorelov). 

In severe winters with plentiful snow, when heavy sleet affects even wild 
and domestic ungulates, the warmth-loving hyaena is not adapted to deep 
snow cover and perishes. At places where hyaenas live near water holes, they 
sometimes die if buried under ice. For example, in the last century in the 
Trans-Caucasus they ' 'often drowned in Lake Karayaz by falling through 
the ice; three bodies of dead hyaenas were found one spring" (Ognev, 1931). 

Population dynamics are not clearly understood. 

Field characteristics. The size of a hyaena is that of a large species 
of dog, for example the German shepherd. The head is massive with broad 
and blunt muzzle and large triangular ears pointed upward. Along the 
47 middorsal neck and back there is a well-developed, erect mane. The anterior 
part of the body is strongly elevated and the rear set low, as a result of which 
the animal seems to rest on its hind legs. The front legs are bent. The short 


tail is equal in length to the thigh and shaggy. From a distance the hyaena 
appears almost white; dark transverse stripes on the flanks and the erect 
mane on the neck are only visible in good light. The animal walks or trots. 
When frightened, it runs in a characteristic heavy gallop, tucking its tail 
between the hind legs, turning its ears back, and exudes a strong unpleasant 

The footprints of the forefeet are considerably larger than the hind ones. 
Undisturbed footprints of the forefeet in a solonchak measure 10 cm in 
length and 8.0 cm in width; corresponding measurements of the hind feet 
were 8.0 and 6.0 cm. When the animal is walking quietly, the footprints of 

46 Fig. 16. Feces of hyaena with characteristic white coloration. Plateau in Kyzyl- 

Dzhar area (see Figs. 10 and 11), Badkhyz preserve. May, 1962. Photograph by 

A. A. Sludskii. 


the hind feet do not coincide with those of the forefeet (as in wolves, jackals, 
and foxes), falling somewhat behind or sideways. At a normal pace the 
distance between imprints of front and hind feet is 55 to 63 cm. Usually, 
impressions of the four claws are visible in all footprints. 

Feces of hyaenas that are feeding on bones are very typical (Fig. 16). 
They are white, in the form of slightly flattened balls, varying in diameter 
from 0.5 to 2.5 cm, and usually consist almost exclusively of disintegrated 
bones. They occur in heaps and are often found in pits around the burrow 
(A.A. Sludskii). 

The voice is a typical cracked howl which can be heard, though rarely, 
at night in all seasons of the year. Many hunters cannot distinguish it. A 
wounded animal growls. (A.S.). 

Practical Significance 

The pelage of the striped hyaena is coarse and sparse; it is not considered 
a fur-bearing animal. Hunting for hyaena is not specially organized; the 
animal is incidentally caught in traps set for other animals. It also dies of 
poisons used in baits for wolves, or is chased and shot. It is caught from 
time to time with the help of hunting dogs and killed later, at a more 
convenient time. As a matter of fact, the hyaena has no fur and hence the 
few skins sold by hunters are generally thought to be wolf or dog skins 
of poor quality. Depending on the quality of the skin and its defects, the 
selling price ranges from 45 kopecks to 1.0 ruble 80 kopecks. Hyaena skins 
are used for preparing chamois leather. 

Many hyaenas were formerly caught in Turkmenia, and about 130 skins 
were collected between 1931 and 1937. Commencing from 1948 in this 
republic, only a few tens of skins were received for curing, and almost none 
received by 1970. In the Trans-Caucasus, Tadzhikistan, and Uzbekistan, 
only a few stray hyaenas were caught in the 1960s and their skins seldom 
sent for curing. For the Soviet Union as a whole, in the 1930s about 200 
skins were processed, less than 100 in the 1950s, and processing actually 
ceased by 1970. 

This animal is of considerable interest to zoological gardens. Live 
hyaenas are mainly caught in Turkmenia and in small numbers in southwest 
Tadzhikistan. Five hyaenas were caught in Tadzhikistan in 1949 and 1950. 

In Turkmenia female hyaenas were earlier highly prized. A belt cut 
from the abdomen together with the genital region, conforming to certain 
patterns, was considered a powerful talisman and was in great demand. 
Magical properties were attributed even to the tail. Dead hyaenas were 
thrown away but the tail always removed (V.G. Heptner). 

The hyaena is little known as an enemy of animal and game husbandry 


in the USSR. In Turkmenia the animal causes damage insofar as it steals 
foxes, cats, hares, and even goitered gazelles from traps, making off with 
the trap (V.G. Heptner). At present, because the hyaena population is small 
over much of the range, the damage these animals cause is negligible. In 
India hyaenas living in the environs of villages cause "considerable damage" 
by destroying goats, sheep, calves, and small dogs (Pocock, 1941); they are 
48 thiLS a ' 'real enemy of livestock." The damage they inflict on melon patches 
by feeding on water- and muskmelons is insignificant. 

In the last century the hyaena had a very bad reputation as a man- 
eater. In the 1880s the newspaper "Kavkaz" reported that some predatory 
animal had attacked human beings, especially sleeping children, for three 
years in Igdyr' Erivansk province; in just one year 25 children were wounded. 
Three adults were also bitten in the same region. Attacks invariably occurred 
at night on people sleeping in the open, in gardens or farms. The hyaena 
was labeled the culprit. The police administration announced a reward of 
about 100 nibles for every hyaena killed. Attacks on man by hyaena were 
reported later from other areas of the Trans -Caucasus, e.g., Surmalinsk 
district of Erivansk province; such attacks were also reported in 1908 
("Okhotnich'ya Gazeta," nos. 25 and 38). Those well acquainted with the 
fauna of Trans-Caucasus (Dinnik, 1914; Satunin, 1915) considered the 
foregoing information reliable and also blamed hyaenas for these deaths. 

Instances of hyaena stealing and killing children sleeping in courtyards 
were reported in Azerbaidzhan even in the 1930s and 1940s (Piraze Agdash 
region, 1939; Mardzhanly Dzhabrail'sk region, 1949). In 1942 in the 
Ismailinsk region (Golyndzhakh) a hyaena entered a hut where the guard 
was sleeping and mauled him (P.P. Aliev). In southeast Turkmenia (Badkhyz) 
hyaenas have reportedly carried away small children during the night (Yu. 
K. Gorelov). The carrying away of a child by a hyaena around Serakhs 
in Turkmenia was reported as late as 1948 (V.G. Heptner). Instances of 
stealing of small children have also been reported in India (Pocock, 1941^. 
Hyaena attacks on sleeping children in Tadzhikistan and Uzbekistan have 
not been confirmed. Sometimes wounded animals attack man (Dinnik, 1914; 
Satunin, 1915). 

Hyaenas sometimes dig up and eat human bodies when the burial is 
shallow and no wooden coffin is used (Berkhnii Guzlak, Fizulinsk region, 
1936; Lembran, Bardinsk region, Azerbaidzhan; F.F. Aliev). 

In ancient Egypt striped hyaenas were domesticated. They were raised 
as livestock on special food. The habit of feeding hyaenas continued in 
North Africa up to the 20th century and was practiced, for example, by 
the Tuaregs (Ronnefeld, 1969). 

At present, over much of the range in the USSR (Trans-Caucasus, 
southwest Turkmenia, Tadzhikistan, and Uzbekistan), the hyaena has become 


a rare animal and causes almost no harm to the national economy. It poses 
very little danger now even for human beings. 

As a rare and extremely interesting animal to scientists, extinction of 
the hyaena should be prevented throughout its range in the Soviet Union. 
In Azerbaidzhan Soviet Socialist Republic the law (1970) baas their hunting 
year-round and a severe fine is imposed for unauthorized killing. Catching 
a hyaena when it attacks man or domestic animals and licensed shooting 
for scientific purposes are permitted. (A.S.) 

Family FELIDAE Gray, 1821 

Felidae are liighly specialized carnivores, in fact the most specialized family 
in the suborder and order. 

Size is small, moderate, or large. The smallest species has a body length 
of about 50 cm and weighs 2.0 to 3.0 kg, while the largest species (Amur 
tiger, Panthera tigris altaica) measures up to 300 cm in length and weighs 
up to 390 kg (ratio of body length between the smallest and the largest 
1 : 6 and weight 1 : 130 to 195). The dimensions of the tiger are inferior 
only to those of brown or polar beat's. All species of the family, even those 
widely differing in external appe<irance (lion and cheetah) have an extremely 
49 characteristic, typical feline appearance, and are essentially monotypic. With 
a fairly large number of species (35), this family is incomparably more 
uniform than the family with the greatest species richness — viverrids 
(Viverridae, about 75 species), or weasels (Mustelidae, about 70 species) 
and even than dogs (Canidae), comprising fewer species (29). 

Felids are of relatively light build, very agile, with elongated (in some 
cases veiy long) tmnks, and usually a long tail. Tail length is not less than one- 
third, and sometimes more than one-half, the body length, and less than one- 
third only exceptionally. The legs are not long, but powerful, and the general 
outline of the body can be described as a parallelogram, sometimes extended. 
In some species (lynx, caracal, and especially cheetah) the legs are long 
and the body profile squarish or nearly squarish. The thorax is small in 
volume and not prominent (lesser volume of lungs and heart); the body 
appears laterally compressed and the thorax is no larger than the abdomen 
in volume. The forelegs are straight while the hind legs in most species 
are somewhat more powerful, although the angles between the foot, shank, 
thigh, and pelvis are relatively small. The sacral region is raised and 
stands above the shoulder region; the posterior portion of the body appears 
particularly powerful, more powerful than the anterior. The dorsal profile 
of these species (a majority) is elevated posterioriy, being sometimes (quite 
rarely) almost straight. Such a particulariy full development of the animal 
body favors a powerful leap from a standing position (without ninning), 


which is typical of all smaller species and some larger ones (puma, snow 
leopard). In a few species (big cats, i.e., lion, tiger, leopard,^ and jaguar), 
the anterior region is more powerful, the back slopes upward anteriorly, 
and the shoulder region stands above the sacral, or more rarely the dorsal 
profile is straight (Fig. 30). 

Neck of moderate length or short. Head relatively small, with a short 
facial portion, and muzzle short and blunt. In some forms (some cats of 
genus Panthera) facial portion relatively large, elongated, and massive, but 
not so extended as in members of the wolf family. In many, the facial portion 
is so shortened that the shape of the head is almost spherical. In all these 
cases the chin region is shifted to the level of the profile of the bare nasal 
region; this is due to the position of the incisors and the position and intense 
development of the canines. 

Eyes large; much larger and protuberant in smaller species; pupil 
rounded or slit. Ears liighly variable in size and form; in some (large species) 
they are not very large, sometimes almost concealed by hair (lion, tiger, 
and snow leopard in winter), but are usually (small species) large, rounded 
or pointed at the tip, set at a slight angle to the sagittal plane, or more rarely 
very highly inclined sideways ("hung out"). The ears, when of a significant 
size, are sometimes set erect, or almost so, pointed, and carry long hair 
at the tip (tuft as in lynx and caracal). In some the ears are set erect, rounded, 
and very large (several species). The total area of the ear opening may be 
more than the area of forward projection of the head. From the outside the 
pinna is slit in the middle and the antitragus well developed, simple, and 
with a cylindrical thickening. The considerable development of the external 
ear corresponds to a significant development of the bony resonators; hence 
the sense of hearing is one of the most important senses of various species 
of the family. Lip ("whisker") and brow vibrissae are well developed, 
particularly the fornier, especially in small species; mandibular vibrissae 
are absent. 

Limbs digitigrade. Forelegs with five digits each, one of which is set 
high and very small. Hind legs with four digits each. Paws of fore- and hind 
50 legs (especially the former) powerfully developed, digits wide-set (paw 
spread out and not compact as in wolves); and digital pads and large 
triangular sole pad well developed. Single small carpal callosity present; 
metatarsal callosities absent. Lower surface of paw usually hairless, and 
callosity in most species well formed; its impression is well seen in pug 

^Throughout this section the terni bars [leopard] is used, which has been reserved from 
early times for the species Panthera pardus. Recently, it has often been written "leopard" 
in the Russian language, a foreign word alien to the Russian tongue. Nowadays bars is being 
employed for Uncia iincia. For this species I have used the term snezhnyi bars [snow leopard] 
or irbis [ounce] (see p. 276). 


marks. However, in some species, especially in winter (lynx) or even year- 
round (sand cat, Felis margarita), hair on lower surface of paw so dense 
and long that it completely conceals the pad. All four digits on each limb 
roughly similar in size; digits III and IV longer but equal in length, and 
II and V equal in size but somewhat shorter than middle ones. On the whole 
cat paws are broad and rounded. All the digits are interconnected with an 
elastic dermal membrane developed to different degrees but, in the majority 
of species, reaching digital pads or claws (see Fig. 278). Digits are extremely 
mobile and can move widely sideways. This is an important feature of the 
cat's paw as a grasping organ. 

Cat digits bear powerful claws. They are relatively short but strongly 
curved, sharply compressed from the sides, and very sharp. The claws in 
all Russian cats, except cheetah, are retracted; i.e., in the resting position, 
and during movement they are shaфly drawn backward and raised. Their 
sharp tips, though pointed generally downward, do not reach the ground. 
Cats move on the pads present at the joint between phalanges I and II. The 
special articular surfaces of the phalanges permit a very broad independent 
vertical movement, while a system of powerful ligament permits retraction 
and extension of their claws (Fig. 17). Two large ligaments extend along 
the top and bottom of all the phalanges, which are firmly attached in front 
to phalanx I; these, in addition to a system of tiny ligaments connecting 
phalanges I and II, play a special role. In the resting position phalanx III 
is set at an angle of about 45° with its distal end turned upward; phalanx 
II in relation to III forms almost a right angle and is set downward. Phalanx 
I is pressed closely to phalanx II. The main axis of the former is parallel 
to that of the latter and raised up and slightly back. In such a position the 
claw is retracted. When the claws are extended (a volitional act) by activation 

Fig. 17. Skeleton and ligamentof digits of tiger showing position and mechanism of 
daw release. Upp)er sketch shows position at rest witli claw retracted; lower sketch shows 
claw extended. Anatomical data from V. Mazak, 1965. Sketch by N.N. Kondakov. 


of the ligaments, mainly the lower one, phalanges III and II align; phalanx 
I, in relation to II, is so positioned that it is vertical and somewhat projected 
forward. The claw is extended but its tip bent downward. The phalanges 
as a group can also bend downward. The horizontal flexibility of the 
phalanges, extension of claws, and downward bend of the digits make a 
cat's paw a powerful and complete grasping device. 

The clasping capacity of a cat's paw is mostly determined by the form 
of its claws. Their curvature varies but is always strong, in several the outer 
margin of the claw forming a smooth arc of small radius. In several species, 
especially in large cats, the claw is firmly attached to the phalanx in such 
a way that its base is enclosed in a well-developed, small bony collar. On 
the whole, the gripping mechanism is better developed in the front paws, 
which also play the role of chief grasping instruments (Fig. 17). 

In a typically well-developed cat's paw each claw has a well-defined, 
highly extensible skin fold which covers the retracted claw laterally and 
dorsally. These "sheaths" protect the claw and its tip. Their ''stretch" when 
51 the claw is released is quite remarkable.^ In species in which such sheaths 
are well developed the interdigital membranes are likewise generally well 
developed and reach the digital pads and claws. In some species, however, 
claw sheaths are wholly or partly reduced, occasionally even poorly 
developed. Interdigital membranes may also be reduced to some extent. 
Both structures are greatly reduced in cheetah (Fig. 278). The paw of this 
animal is characterized by other features — claws less sunken and externally 
visible (see characteristics of cheetah). 

Claws come into play only during gripping, holding, or killing prey, 
or in self-defense, and to a lesser extent when climbing trees; they are not 
iLsed at all in locomotion or for digging. Thus their tips are always sharp 
and ready for action. At the same time, the anatomy of the paw is probably 
one reason why cats cannot run fast or long. Only cheetah are fleet-footed 
and hunt prey by sprinting. Among mammals, they are the swiftest animals. 

A paw with retractile claws is a characteristic feature of the family; 
among other carnivores, this feature is seen only in a small group of viverrids 
(Prionodontinae; genera Prionodon and Pardictis). 

In most animals the pelage is dense, soft and silky. The length, 
compactness, and extent of growth of underfur vary considerably, depending 
on climatic conditions in the habitat. In severe climates seasonal differences 
in coat color are distinct. The length of hair throughout the body is fairly 
uniform in most species. Only in a few (lynx) are long "side-whiskers" 
found on the cheeks. In male lions a dense, long mane occurs on the anterior 

■'Some references in literature to the presence of horny caps at the base of claws 
(Sapozhenkov, 1957; and others) are incorrect. They are evidently mistaken descriptions 
of the dennal folds described above. 


part of the tmnk, neck, and head up to the ears; long hair also occurs along 
the chest and abdomen. In cheetah elongated hair likewise occurs along 
the nape and on the withers. 

Coloring extremely variable, but mostly spotted, with dark spots lying 
on a fundamentally light background. The shape of the spots varies greatly, 
from simple, close-set, small dots, to large ring-shaped areas consisting 
of individual spots, completely circular or elongated rings, and so on. Quite 
often the spots form fairiy regular longitudinal or transverse rows; some 
merge into transverse or longitudinal stripes; coloration is thus a combination 
of strips and stretched and circular spots. The tnmk region of some species 
is devoid of spots, consisting only of vivid transverse stripes (tiger). 

The general background color is usually a light shade, varying from 
almost white or light yellow to rusty, bright rusty, or cinnamon. Spots are 
usually black, of varying intensity and shaфness of outline, and sometimes 
nist-tinged. Some species are uniform in color (caracal, puma and lion) 
or almost so (sand cat, Fells margarita); their color is usually sand-yellow, 
brownish-yellow, or some similar shade. Sometimes against a uniform 
general background very poorly developed spottedness is visible, or only 
traces of it on the limbs (some forms of lynx), or a pattern on the head 
(one of the colored forms of Temminck's cat, Fells temmlnckli). Usually, 
even among monochromatic species kittens or juveniles have more or less 
developed spots. The spotted pattern of present-day cats should be considered 
as an ancestral character. 

Individual color variability is pronounced in many species, as is 
geographic variability. Geographic variability is generally exhibited in a 
change in overall color shade, and often in type of spottedness. In some 
species melanism, albinism, and chromism are encountered, while geographic 
localization of such abnormalities is known [black ''panthers" (leopards) 
of India, chromic manuls of Turkmenia, and others]. Some species show 
52 seasonal variation in color {Fells yaguarundi of America). In one case (Fells 
temmlnckli) the color, although not being in the strict sense polymorphic 
changes very 8Ьаф1у (spotted to uniform, dark to light); geographic 
localization of color types is also known. Among lions sexual dimoфhism 
is manifested in the growth of a mane in males. In all cats males are larger 
than females. 

Anal glands are usually present but poorly developed. Penis located 
directly in front of the scrotum. Glans rudimentary and with homy spines. 
Teats, two to four pairs. 

The skull is characterized by several distinct family features. However, 
as a result of significant differences in the absolute overall size and other 
characteristics of different species, the general appearance and skull 
proportions may vary considerably. In general features the skull is broad. 


with widely diverging zygomatic arches. Anterior part of skull, from anterior 
margin of orbit ("muzzle"), short and broad. Ethmoturbinal scrolls large, 
extend forward almost up to opening of nasal chamber, and lie on top of 
maxilloturbinals. Alisphenoid canal absent. 

Bony auditory bulla usually high, rounded (swollen), formed of 
ectotympanicum and entotympanicum, and divided internally into two 
chambers by a septum. Entotympanic (rear-medial) portion invariably, but 
to varying degrees, larger than ectotympanic portion, which communicates 
with ear opening. External separation of two portions of bulla not usually 
perceptible and distinct only in some forms. Auditory tube absent or barely 
discernible. Relative size of tympanic bulla highly variable; large, swollen, 
and thin-walled in small species (especially in desert species); relatively 
small in large animals and often flat and thick-walled. Mastoid and 
paroccipital processes separated; latter flat, usually closely adhering to 
posterior wall of bulla, and in a majority of species does not project beyond 
lower margin of tympanic bulla. 

The following are general changes in the overall appearance of the 
skull. Skull of more or less neutral type (lynx, caracal; Vereshchagin, 1967): 
general outlines fairly elongated-oval; cross section about 75% of 
condylobasal length. Interorbital and postorbital constrictions broad, cranium 
very voluminous, weakly extended forward, and highly enlarged; zygomatic 
arches widest at level of articular surfaces but their general outline bulges 
or is arched. Crests and tuberosities for attachment of muscles, especially 
masticatory ones, weakly developed; sagittal crest present but small, while 
lambdoidal crest well developed. 

The series comprising species of larger and largest cats and culminating 
in tiger is characterized by the skull being very heavy and massive, the 
facial region, especially the muzzle, relatively elongated, and the cranial 
region relatively short. The entire skull assume-s a highly extended form 
although the rear portions of the zygomatic arches may be very broad. 
Viewed from the front the features are not so bulging but appear very angular. 
The interorbital and especially the postorbital constrictions become relatively 
small and volume of the brain case is reduced and not inflated. Tuberosities 
and crests very powerfully developed; sagittal and lambdoidal crests 
particularly large and high; brain case greatly compressed by powerful 
masticatory muscles. Teeth, especially canines, powerful with a broad base, 
but relatively short. Tympanic bullae relatively small, flattened, and thick- 
walled in extreme cases. Lower jaw heavy, relatively long, with a powerful 
anterior portion projecting forward, bearing large canines with long and 
massive bases. This is a skull typical of animals which kill and tear apart 
large quarry. Often the quarry is larger than the predator in size and weight; 
in the very largest by several times. 


53 The skull of small species is characterized, on the contrary, by lightness, 

a thin-walled brain case, and absence of any crests or tuberosities, apart 
from the lambdoidal. Sagittal crest not developed, or represented only by 
a small ridge at occipital end and lyrate figure [temporal crest] usually broad. 
Facial region of skull abbreviated to a greater degree, muzzle short and 
blunt, and orbits highly enlarged and increasingly set forward. Zygomatic 
arches very thin in midportion (not only from the rear) and form a fairly 
regular arcuate shape. What is important is that the brain case, sometimes 
moderately enlarged and slightly extended, becomes increasingly 
voluminous, bulging, and short. Thus the entire skull is short and swollen 
with broad interorbital and postorbital regions (see Fig. 19). 

The general contour of the skull in some forms is roughly oval, as in 
lynx, or even somewhat elongated (elongated brain case), or tends to become 
broader and shorter, and in extreme cases (manul, F. manul) even spherical. 
On the whole the skull of small cats, compared with that of large ones, 
appears infantile. The teeth are relatively weak and the canines relatively 
long, but thinner. The lower jaw is short and light, with a relatively weak, 
not massive, anterior portion. This is the line of specialization of small 
predators which kill small prey, much smaller in size (rodents) than the 
predator itself. If cats are regarded as perfect predators the above two forms 
are superior types, even though morphologically they contrast with each 

Dentition shows extreme specialization within the order with regard 
to number of teeth and certain features of their structure. Complete dental 

.31 3 1 

1 , с , pm , m = 30, 

3 1 2 1 

but in some cases second upper premolar (first in row since first upper 
premolar generally absent in cats) sometimes does not grow (or is lost), 
or is invariably absent (manul and lynx). Dentition in these cases: 

3 1 2 1 
i , с , pm , m = 28. 

3 1 2 1 

The diastema, especially in the lower jaw, is large in most species. 
This is associated with the absence of the first premolar in the upper jaw 
and the first and second premolar in the lower jaw, which in turn is due 
to the intense development of canines and jaw abbreviation. 

The teeth are powerful and sectorial (cutting). Carnassial teeth large, 
highly compressed, and relatively long; incisors small and weak, form 
straight transverse rows, and canines large. In large species they are more 


massive and short and in small species relatively thin and long, sometimes 
very long and thin (clouded leopard), and usually with shallow longitudinal 
grooves. Second premolar (upper) very small, conical and single- rooted; 
third upper premolar well developed, compressed from the sides, usually 
with a high and sharp central cusp, and two small cusps located at its base 
at front and rear. Upper camassial tooth (fourth premolar) very large, 
compressed from the sides, with three cusps lying in a single plane; first 
cusp moderate and conical; second high and sharp; and third low and long 
with a cutting edge. Antero-internal surface of teeth, in most, with additional 
cusp with a small, usually well-developed blunt cusp. First upper molar 
rudimentary, very small, and set across the row (sometimes lost). 

In the lower jaw both premolars (third and fourth) have three cusps, 
like the third premolar in the upper jaw. Carnassial tooth (first molar) 
powerful with two well-developed cutting edges. Milk carnassial in some 
species complex compared with permanent tooth. 
55 A characteristic feature of the skeleton is the great flexibility of the 

spine, and the increased area and special form of the articular surfaces of 
the phalanges and their position in relation to each other, which allows for 
retraction and extension of the claws (see above). Thoracic vertebrae, 13. 
Os penis rudimentary. 

The relative length of the intestine, as a result of feeding exclusively 
on meat, is small; the relative weight of the heart is small, much smaller 
than in species of the dog family. 

Felids live in very diverse habitats, ranging from humid tropical forests 
to sandy deserts. They are usually associated to some degree with forest 
and shrub vegetation (for cover). They inhabit warm or moderately warm 
climates without a permanent or with a slight snow cover. Only one species 
(lynx) is regularly associated with areas of snowy winters, either on the 
plains or in mountains, and another (snow leopard) with high snowy 
mountains. Part of the range of some species includes regions with a snowy 
winter (puma, Amur cat, forest wild cat, tiger, and others). 

All species are predators, mainly of homoiothermal animals of extremely 
varying size, from small rodents and birds to giraffes and buffaloes. More 
rarely, they catch reptiles; some are specialized for catching fish. They hunt 
by lying in ambush along tracks, near water holes and solonetzes, or by 
concealing themselves and overtaking the quarry in one or more leaps. They 
are not capable of prolonged running or chasing. Only cheetah can pursue 
its quarry for some hundreds of meters and overtake even a gazelle, by 
attaining a speed maximum for mammals.-"^ They are solitary animals and 

■^Cheetah are usually considered capable of attaining a speed of up to 140 km per hr. 
According to recent data the maximum confirmed speed is 56 miles (90.12 km) per hr 
(PoumelL 1964). 


do not form pairs even at the time of raising the young. Evidently only 
one species (lion) lives in pairs and family groups and hunts in groups. 
The den is in the open or located in burrows, rock crevices, and caves. 
Cats are predominantly nocturnal animals. 

The range of the family is extremely extensive and covers all 
zoogeographic regions except Australia (Fig. 18). In the New World cats 
are distributed from the northern coast of the [North American] mainland 
to Patagonia and the Strait of Magellan. The family is absent on the American 
Islands in Bering Sea, except Nunivak (?) [sic]. In the Arctic Archipelago 
the boundary of the range encompasses Baffin Island, Southampton Island, 
the small Hudson Bay Islands, Newfoundland, Islands of St. Lawrence Strait, 
and islands along the west coast of North America (Kodiak, Vancouver, 
Alexander Archipelago, and others). Representatives of this family are absent 
in the West Indies, including the Bahamas, Cuba, and Trinidad. On the 
coast of South America, Chile falls within their range. Cats are absent in 
Tierra del Fuego and the Falkland Islands. 

In the Old World the range encompasses almost the whole of Europe 
in the north, to the tree line. In the west the range includes England; members 
of the family are absent in Ireland and evidently have never been present, 
at least not in the historic period. In the Mediterranean Sea the range covers 
Corsica, Sardinia, Sicily, Crete, and Balearic Islands (in the rest of the 
islands, including Rhodes, Cyprus, and Malta, cats are absent). Members 
of the family are absent and were never present in islands of the Barents 
Sea and small islands and archipelagos of the Atlantic Ocean. 

In Asia the northern boundary of the range runs along the northern 
tree line, or a bit to the south, including Kamchatka in northeast Siberia. 
The southern boundary encompasses the Arabian Peninsula, India, Ceylon 
[Sri Lanka], and running along the coast of Bay of Bengal descends south 
to encompass Sumatra and Java, extending between Bali and Lombok, 
encompassing Kalimantan (Borneo), Palawan and some of the Philippine 
Islands (evidently only Negros and Cebu in the center). In the Mentawai 
Archipelago (west of Sumatra), and also on the islands of Billiton, Bangka, 
and Madura (Java Sea), members of the family are absent. Cats are also 
absent in the small islands south of Singapore, between Sumatra and 
Kalimantan (Borneo), and in the South China Sea. Farther, the range bound- 
56 ary encompasses Hainan and Taiwan, leaving out the Ryukyu and Japanese 
islands, but includes Tsushima and Cheju Do (Quelpart), encompassing 
Sakhalin but leaving out the Kurile Islands, and continues toward Kamchatka. 
Members of the family are absent on Karaginsk and the Commander Islands, 
and evidently also on the Shanter Islands. 

The range covers all the African mainland; Madagascar, and Atlantic 
and Indian Ocean islands are excluded. Cats, especially the large ones, have 



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vanished at some places and the range of the family has shnink. On the 
other hand, domestic cats which become feral under certain circumstances, 
especially in tropical countries, have settled on some islands outside the 
natural range of the family. Thus Felis megalotis, described as a separate 
species from Timor (and Roti), in spite of some persistent objections (Weber, 
1928), is considered feral.'* 

The cat family, excludmg the Malagassy Fossa (Viverridae), is a highly 
compact natural group. This family is well separated from related families, 
above all the closely allied civets (Viverridae).'' This is true of the family 
as a whole, i.e., including fossil forms. Even such extreme forms as saber- 
toothed cats are typical of the family. Attempts to raise some fossil groups, 
mainly saber-toothed cats, to the status of families (an extreme point of 
view — Kretzoi, 1929), have not been recognized (Simpson, 1945; Gromova, 
1962). Nevertheless, some recent taxonomists have placed saber-toothed 
cats in their own family (Machairodontidae) (Thenius and Hofer, 1960). 
Proposals for segregating cheetah and lynx from Recent Felidae into a 
separate family (Guepardidae Gray, 1869; and Lyncidae Schulze, 1900) 
are not well justified and have found little support. 

At present it appears more natural to view the family broadly and accept 
five subfamilies: extinct Proailurinae Zittel, 1893; Nimravinae Trouessart, 
1885; Machairodontinae Gill, 1872; Hyaenaelurinae Pilgrim, 1932; and con- 
temporary Felinae Trouessart, 1885. Machairodontinae (Lower Oligocene- 
Pleistocene; about 10 genera) are considered a very specialized group, standing 
in rank above the Felinae (Simpson, 1945). The more primitive cats of the 
subfamily Proailurinae (about five genera) existed from Late Eocene (Early 
Oligocene) through the Middle Pliocene in the Old World. Some of their 
early forms combine certain features of cats and civets ("structural ancestoi^"), 
indicating the course of development of the family and its affinities. 

The original forms ("ancestors") of present-day cats are recognizable 
among members of subfamily Nimravinae (Upper Eocene-Lower Pliocene of 
Eurasia, North America, and Africa; about nine genera). Fossil forms of the 
contemporary subfamily Felinae are known from Asia and in the Pleistocene 

''Tliis form is neither mentioned in the most recent catalogs of the family nor in the 
faunal compilations (Laurie and Hill, 1954, for example). The zoogeographically exceptional 
case related to the possible occurrence of species in the family in the Australian zoogeographic 
region has not been discussed. 

^Its affinity with civets is plainly evident, however. Civets of genera Prionodon and 
Pardictis (linsangs of southeast Asia) have a short penis, characteristic of cats, which adjoins 
the scrotum; they have no perineal glands and the claws are retractile and covered with a 
dermal sheath, another characteristic of cats. 

The isolated position of family Felidae is emphasized by the fact that cats differ sharply 
in serological characteristics from other predators. The range of these differences is greater 
than differences between some carnivores and all pinnipeds (V.I. Borisov). 


(perhaps Pliocene) of North and South America. Apart from various species 
of the genus Fells, two fossil Asiatic Lower Pliocene genera (Slvaelurus 
Pilgr. and Dlnofells Zdansky) and one Pleistocene genus {Slvapanthera 
ICretzoi) have been described. 

Certain aspects of the taxonomy of present-day cats are far from simple, 
and confusion and complications persist even today. This applies to supra- 
57 specific groups (the question of the species of cats is now clear). At present, 
there is hardly a family of present-day mammals in which several conflicting 
viewpoints relating to their systematics cannot be adduced. Thus the 
classification of present-day cats into two subfamilies — cheetah 
(Acinonychinae) and the remaining species (Felinae) — long supported, was 
replaced by three subfamilies by separating Pantherinae, or "big" cats, 
from Felinae (Pocock, 1917; Ognev, 1935; and many others). A later 
suggestion classified present-day cats into four subfamilies by further 
separating lynx (Lyncinae) from true cats (Felinae) (Haltenorth, 1957; 
Weigel, 1961; Mazak, 1965; and others). An extreme view, which however 
evoked no support, proposed the classification of cats into 15 subfamilies. 
A fifth subfamily consisting only of the clouded leopard Neofelinae with 
one genus and species has also been proposed (Kretzoi, 1929). 

The views of various present-day taxonomists on the classification of 
cats at the level of subfamily are highly diverse. Most propose two (cheetah, 
Acinonychinae, and all other cats, Felinae) or three (cheetah, big cats, 
Pantherinae, and the remainder). Some recognize four subfamilies, 
establishing a separate subfamily for lynx (Lyncinae). The position of 
individual species thereby becomes extremely diverse: clouded leopard (F. 
nebulosa) appears sometimes under Pantherinae and sometimes outside this 
group; manul is sometimes placed among Felinae and stands close to sand 
cat {F. margarka) or sometimes in the lynx subfamily (Lyncinae). 

The latter case exemplifies well a formal approach to taxonomy, 
dismpting its naturalness. In all craniological and other characteristics the 
manul is a close relative of the sand cat (see below in systematic section) 
and represents an extreme degree of specialization among "small cats" 
in particular and contemporary cats in general. However, the manul lacks 
the anterior upper premolar (p-) and, based on this single formal 
characteristic, it falls into the nearest genus, in the same group with lynxes, 
with which the manul has hardly anything in common. 

The classification of subfamilies in other cases is even less satisfactory. 
Thus, a comparison of the diagnostic features of the subfamilies of small 
cats (Fehnae), big cats (Pantherinae), and lynx (Lyncinae) (see for example, 
Mazak, 1965) reveals that, in the ultimate analysis, the only diagnostic feature 
of Lyncinae is the tufts on the ears. Quite evidently, this feature is not 
adequate enough for its separation. 


The inclusion of big cats, i.e., lion, tiger, leopard, and jaguar, usually 
also the snow leopard, and clouded leopard (Neofelisy in a separate 
subfamily (Pantherinae), which at first appears quite natural, is also 
unacceptable. There are no shaф differences between this group and Felinae. 
Insofar as the clouded leopard is concerned, it has no features which are 
characteristic of Pantherinae proper, i.e., lion, tiger, leopard, and jaguar, 
while the snow leopard exhibits features of both "Pantherinae" and 
"Felinae" in an almost equal ratio (Hammer, 1967; see also characteristics 
of the genus below). Thus there is no distinct demarcation even between 
typical big cats ("Pantherinae" — lion, leopard, jaguar, and tiger) and small 
cats ("Felinae"), at the subfamily level. Tlie snow leopard is a connecting 
link between these groups; it is customary to consider its features inter- 
mediate (see description). Sometimes contemporary cats are not classified 
into subfamilies; instead all are placed in one subfamily (Felinae) as opposed 
to the extinct species (Ellerman and Morrison-Scott, 1951 and 1966; 
Ellerman, Morrison-Scott and Hayman, 1953). This viewpoint reflects more 
correctly the natural relations of the species of the family but is hardly a 
prevalent one. It is also my opinion that a clear-cut separation of even genera 
58 of contemporary cats is not only difficult but almost impossible (see 
characteristics of genera below). This is true of even the cheetah; the features 
of which always call attention to it. The actual differences in this genus 
are fewer than generally considered (see characteristics of the genus below). 
Thus, only one subfamily can be identified among contemporary cats.^ 

The situation in the generic groupings of present-day cats is chaotic. 
This is due not so much to the complexity of the problem itself, as confusion 
in the concept of genus in modem taxonomy, primarily due to the aversion of 
some specialists in systematics for large genera and their attempts to create 
genera with fairly equal numbers of species.^ Thus a general tendency to 

*In spite of its large size, the puma exliibits no affinities with the group of big cats 
{genus Panthera, "Pantherinae"), a fact long recognized even by advocates holding extremely 
disparate views. Although this animal is large, it is a true small cat (Felis) in all moфhological 
features. Herein we do not place puma among big cats (whatever be the inteфretation of 
the group) and have no intention of doing so at any time. 

^The number of works devoted to the supraspecific taxonomy of cats is very large. The 
work of N A. Severtsov (1858) is the foremost and most interesting study; it exerted great 
influence on all subsequent works and enjoys importance even today. The works of Pocock 
(1917) and Haltenorth (1936 and 1937) are also significant. At present, several works have 
been published which are essentially moфhological; even ethological works have been 
published in rex^ent years (Weber, 1928; Kretzoi, 1929; Tsarapkin, 1932; Ognev, 1935; Pocock, 
1939 and 1951; Simpson, 1945; Leyhausen, 1950 and 1956; Haltenorth, 1953; Thenius and 
Hofer, 1960; Weigel, 1961; Hemmer, 1964; Mazak, 1965; Vereshchagin, 1%7; and others). 
Much of the material on big cats has been reviewed and revised by Hemmer (1966). 

^See Heptner (1965) for "larger" genera in the structure of systematic groups and 
their evolutionary significance. 


further divide genera has developed. Cats, in this respect, provide excellent 
material and there is no other group of mammals in which this tendency 
has been more manifested, resulting in many major contradictions and 
distortions. Yet there are some contemporary taxonomists who project a 
fairly natural picture of supraspecific groupings of cats. All of them propose 
a very small number of genera. 

In essence, N.A. Severtsov (1858) was the first to make a broad, 
theoretically based analysis of the supraspecific groupings in the family. 
He established a total of 5 genera with 27 subgenera: Tigris (lion and tiger), 
Pant/iera (jaguar, leopard, ounce, and puma — four subgenera), Cynailurus 
(cheetah), Lynchus (lynxes and caracal — two subgenera), and Felis (all 
remaining cats — 17 subgenera). Severtsov 's system, which by and large 
is well substantiated and thought through, remains unsurpassed even today, 
Pocock (1917) considered Severtsov's classification basic, from one of the 
modem viewpoints. However, Severtsov's concepts were altered greatly in 
principle by Pocock in that most of the subgenera proposed by the former 
author, primarily in the genus Felis, were raised to the rank of genus. Thus 
Severtsov's natural genus Felis, which is "large" by modern standards was 
downgraded. Pocock (1917) divided the family into 17 genera and subgenera. 

Some contemporary theoreticians have split these genera arbitrarily 
or follow approximately the same combination. Thus, some accept (while 
maintaining 3 or 5 subfamilies) 14 (Haltenorth, 1957), 19 (Weigel, 1961), 
20 (Mazak, 1965), and even 24 (Ognev, 1935) genera. This system of 
classification is dubious even on theoretical grounds. According to Mazak 
(1965), of the 20 genera, 13 (65%) are monotypic, 1 contains 2 species, 
3 contain 3 species each, 1 contains 4 species, and another 5 species. With 
24 genera (Ognev, 1935), the number of monotypic genera rises even higher. 
Thus generic characteristics, features, and the very concept per se become 
species-based. The natural concept of a monotypic genus in such a case 
is largely lost. 

In point of fact it appears at times that between some species within 
the same genus differences are incomparably larger and more significant 
(Mazak, 1965) than differences between genera {Felis — Prionailurus; 
Lynx — Caracal; and so on). Such a situation is arbitrary and difficult to 
comprehend. Various authors not only evaluate differently the rank of 
individual forms (big cats — lion, tiger, leopard, and jaguar — one genus 
or four monotypic genera, and so on), but they also occupy very different 
positions in the series (evolutionary, specialization) of forms in the family. 
Thus, the manul is placed as a distinctive genus sometimes in the immediate 
59 proximity of genus Felis (Mazak, 1965), sometimes there are 14 genera 
and 7 subgenera intervening between them and it is placed in a separate 
"subfamily," i.e., lynx (Weigel, 1961), and so on. All this underscores the 


artificiality of excessive subdivision of contemporary members of the family 
into genera. 

An opposite, extreme viewpoint recognizes only two genera — Acinonyx 
(cheetah) on the one side and Fells, which includes all remaining 
contemporary cats, on the other. Usually the latter genus is extensively 
subdivided into subgenera. This system, which is quite reasonable (see later), 
has several supporters (Allen, 1938; Novikov, 1956; Hall and Kelson, 1959; 
and others). 

Yet in spite of all the diversity of size and shape of individual species, 
large craniological differences, and relatively large number of species (35), 
contemporary cats represent an extremely homogeneous group moфhologi- 
cally, systematically, and phylogenetically. In practice, the position is that 
almost any two species which are extremely dissimilar can be linked through 
transitional forms into a fairly continuous moфhological series. Breaking 
this chain or mosaic pattern into innumerable independent genera is almost 
invariably artificial. Even the characteristics of the cheetah (see below) are 
less 8Ьаф and more typical than usually admitted. This applies to behavioral 
characteristics also (Leyhausen, 1950 and 1956; Hemmer, 1964 and 1966; 
and others). 

It is interesting that in [almost] all species, as far as is known, the number 
of chromosomes is identical (2n = 38) and the differences in karyotype 
insignificant; only two {F.pardalls and F. wledi) have 36 chromosomes* 
(V.N. Orlov). Under artificial conditions (zoological gardens) all species 
of large cats have produced hybrids (lion, tiger, leopard, and jaguar) and 
even trihybrids; backcrosses are also successful. So far only in the case 
of the snow leopard are hybrids not known; however, it does not breed 
readily in captivity; and tiger with leopard (information relating to such 
hybrids in nature is not reliable). But then, hybrids have been produced 
by crossing leopard with puma, which is not a big cat (genus Panthera), 
i.e., a case of intergenus hybrid (Hemmer, 1966). 

The diversity of interpretation of the above-noted characteristics of the 
family notwithstanding, natural groups are known which, according to some 
other possible grouping of the species, could be assigned the rank of genus 
among present-day cats. Three or four such groups are recognized. According 
to one point of view (Simpson, 1945), the family consists of three genera — 
big cats, Panthera (lion, leopard, tiger, jaguar, snow leopard, and clouded 
leopard), cheetah (Acinonyx) and small cats (Fells) including all the remain- 
ing species of the family. Here the subfamilies of Pocock (1917) are treated 
as genera. According to an identical viewpoint (Ellerman and Morrison- 
Scott, 1951 and 1966) the same genera with similar species composition 

*As of time of publication of Russian edition — Sci. Ed. 


are recognized; however, the clouded leopard (F. nebulosd) is allocated 
to an independent genus (Neofelis), occupying a position between the genera 
Panthera and Felis\ in all four genera are recognized (Palearctic and 

It is becoming increasingly clear (see the new works cited above, 
especially that of Hemmer, 1964 and 1966) that Neofelis has no affinities 
with big cats {Panthera). Neither does it assume an intermediate position 
between big and small cats (Felis). This is, apparently, a genuine small 
cat which is evidently no more peculiar than some other more or less 
"peripheral" species of this group. It is placed here as a member of genus 
Felis (as a subgenus). On the other hand the snow leopard is not wholly 
typical of the genus of big cats (Panthera) and combines fairly equally the 
characteristics of both big (Panthera) and small (Felis) cats.^ However, it 
60 differs from both, possessing features typically its own, and, in spite of 
also exhibiting features of an intermediate nature, is separated as an 
independent genus. ^° 

Thus, as in the works of other researchers cited above, four genera 
are recognized here also, but somewhat differently and with differences 
in species composition: big cats (Panthera Oken, 1816), ounce or snow 
leopard (Uncia Gray, 1854), small cats (Felis Linnaeus, 1758), and cheetah 
or pardus (Acinonyx Brookes, 1828).^^ 

The genus cheetah (Acinonyx) is unique and well separated, albeit less 
well so than commonly admitted. This genus is close to small cats (Felis) 
and is undoubtedly derived from them. The cheetah is a specialized cat, 
the characteristics of which he in adaptation to fast running, i.e., in a direction 
altogether unusual for cats, and to catching relatively large prey. It exhibits 
no affinities with big cats (Panthera). 

The demarcation between genera of big cats (Panthera) and small cats 
(Felis) are far less definite. Significant features of an intermediate character, 
as pointed out above, are seen in the snow leopard (genus Uncia), which 
connect them quite clearly. There are also forms of trae small cats exhibiting 
affinities toward the snow leopard (clouded leopard, puma). Evidently these 
are features of convergence resulting from the large size of the animal as 

^lis situation may provide a base for combining genera of big (Panthera) and small 
(Fslis) cats in a common genus. This approach is possible; however, it applies only to those 
subgeneric groups which could be separated within the limits of genera Felis and Panthera, 
in relation to the taxonomic weight of their features. 

^''Details of all genera are given later under their description. 

^'With reference to the degree of taxonomic fragmentation of contemporary cats and 
in the approach to supraspecific groupings in the family, i.e., few genera and extensive use 
of the concept of subgenus, the views of some contemporary scientists (Ellemian and 
Morrison-Scott, Simpson, and Heptner) come very close to those of N.A. Severtsov (1838). 


well as its prey. Thus Panthera, Uncia, and Felis form a close group or 
a chain of successive genera. 

Relative to the level of organization ("primitiveness" or "progressive- 
ness") of these genera and the degree of their moфhological specialization, 
it is quite difficult to resolve the question of monotypic species in the family 
and the closeness of their relationships are quite troublesome. One is 
hampered additionally because paleontology provides very little data due 
to the fact that various types of present-day cats appeared quite 
simultaneously. Evidently even the directions of various adaptations among 
present-day Felinae also proceeded more or less simultaneously. According 
to the traditional and more prevalent point of view, all small cats of the 
genus Felis form a least specialized ("primitive") group, and big cats of 
the genera Panthera and Uncia, as well as cheetah, a more specialized 
("progressive") group. The latter is usually considered specialized in a 
special sense and in a direction different and independent of big cats. 

In giving a very general evaluation of the skull structure of small cats 
of the genus Felis (apart from those such as puma and clouded leopard), 
attention is drawn to features such as the very large volume and bulging 
form of the cranium, the very short facial region, and the absence or slight 
development of various types of crests and tuberosities serving for the 
attachment of muscles, mainly the group of masticatory muscles. These 
features are usually considered juvenile (infantile) and often justifiably 

Fig. 19. Skull of tiger {Panthera tigris L.) and manul (Felis manul Pall.) reduced 

to the same scale, two extreme types of contemporary cats. Collection of the 

Zoological Museum, Moscow University. Sketch by N.N. Kondakov. 


regarded as an index of the slight specialization of the group, that is to 
say, more neutral than those of more primitive structure. The opposite type 
of features characterizing skulls of big cats are considered specialized and 
hence progressive. 

This traditional approach is hardly applicable to contemporary cats. 
The skull of big cats with its highly developed facial region, powerful crests, 
and relatively moderate, elongated braincase, has an appearance and structure 
of the skull far more similar to that of predators of other families than does 
the skull of small cats. The skull of big cats exhibits a greater degree of 
general predatory features than specific feline characters and structure 
61 (Fig. 19). Members of the small cat genus Felis s. 1. exhibit features that 
are more sharply specific for contemporary cats. The differential 
characteristics of modem Felidae are more fully manifested in the skull 
of small cats. Thus progressive specialization within present-day cats 
proceeded in the direction of the development of a superior feline type, 
as seen in the manul but not in tiger. In spite of all the specialization of 
big cats, typical feline characteristics in them are fewer than in small ones 
and less distinctly manifested. Small cats are tmer representatives of cats 
in the total meaning of the term. 

In spite of scanty paleontological material, one can conclude that 
present-day big cats (genus Panthera) exhibit a larger number of features 
of similarity with fossil forms, even with totally extinct groups, than present- 
day small cats. Fossil forms characteristic of the Felis type {silvestris, for 
example, or lyjix), not considering the most recent direct antecedents to 
contemporary forms, are either unknown or few in number. Evidently the 
evolution of a fully developed type of contemporary small cat (genus Fells) 
represents a new stage in the evolution of the family and subfamily. The 
relative primitive nature of big cats has in fact been noted by paleontologists 
though, it is true without arguments (Thenius and Hofer, 1960), although 
usually they support a contrary view (Simpson, 1945). 

The placement of modem cats with a ''juvenile" type of skull with 
progressive members of the family is unusual, but the situation is not unique 
in predatory mammals (Ferae). The family of common seals (Phocidae), 
invariably recognized as the most specialized and progressive in the order 
(suborder), differs from eared seals (Otariidae) in skull structure. For 
example, the genus Eiimetopias (sea lion) exhibits particulariy well- 
manifested overall predatory features in the skull. The course of development 
of the family evidently lay in the course of cephalization. 

Increase in overall size in a phylogenetic series is usually considered 
a sign of specialization and advancement. It is, however, hardly possible 
to assign an absolute and decisive importance to increase in size; size is 
extremely relative in these same pinnipeds. The number of species in the 


genera also underscores the progressiveness of the small cat group (Heptner, 
62 1965): about 11.5% of the species in the family fall in the genus of big 
cats (Panthera), 3.5% each in the genera of cheetah (Acifionyx) and snow 
leopard (Uncia), and 81.5% in the genus of small cats. 

Thus there is adequate justification to begin the phylogenetic series 
of contemporary cats with big cats (Panthera), followed by snow leopard 
(Uncia), and then small cats {Fells). The cheetah {Acinonyx) represents a 
specialized lateral branch of the line of small cats. These groups represent 
three main directions of specialization with respect to size of prey and method 
of hunting — catching large animals (big cats and snow leopard), catching 
small vertebrates (sometimes larger; small cats), and catching large animals 
by pursuit (cheetah). 

The family Felidae contains fewer species than civets (Viverridae, about 
75 species) and weasels (Mustelidae, about 70 species); nevertheless with 
35 species the family comprises 14.5% of the total number of species in 
the order and about 1.0% of all mammalian species. Felidae outnumbers 
the dog family species-wise (Canidae, 29 species). In the genus of big cats 
{Panthera) there are four species; ounce or snow leopard (Uncia), one; small 
cats {Fells), 28; and cheetah {Acinonyx), one. 

Of the four genera, one (cheetah) is Afro-Asian; one {Panthera) mainly 
Old World with one representative (jaguar) in South, Central, and North 
America (extreme southern part of North America*); one {Uncia) wholly 
Asian; and one {Fells) widely distributed throughout the range of the family 
in the Old as well as New World. The maximum number of species are 
associated with the Old Worid, particulariy Asia; the smallest number inhabit 
North America; a fairly significant number are found in South America. 

The economic importance of the family as a whole is minimal and 
depends on the level of economy in a given region. In untouched biocenoses 
and regions little settled by man, species of the family have little or no 
practical importance as fur animals or in the trade of live animals for 
zoological gardens. In more densely populated regions big cats often kill 
cattle or damage game prospects by killing wild ungulates. Some individual 
animals (tiger, lion, sometimes leopard and jaguar) pose a danger for man 
under certain conditions in a given region. 

Some large species (lion, tiger, leopard, puma, and jaguar) are valued 
as game and sport animals. At the same time they exert a selective influence 
on wild ungulated populations, which is a positive factor. On the other 
hand, in cultivated areas even small cats may damage the hunting economy 
by destroying game birds (pheasants), hares, or young ungulates. Many 
species feed almost exclusively on small rodents but are seldom effective in 

*Closing parenthesis out of place in Russian original — Sci. Ed. 


controlling rodent populations. Some species are protected in certain areas 
or sanctuaries. One (F. silvestris) also survives as a truly domesticated animal 
(domestic cat, "F. catus L.")^^ and two species have been tamed and 
trained for hunting (cheetah for hunting gazelle, and caracal for hunting 
birds); but cannot be considered "domesticated." 

In all, 4 genera and 11 species, i.e., about 31 to 32% of species of 
family Felidae, are found in the Soviet Union. They comprise about 10% 
of the species of the Russian fauna. 

The range of Felidae covers almost all of the northern USSR to the 
tree line. The majority of species are associated with the southern regions 
of the country, however; only one (lynx) also occupies extensive regions 
in the north. There are forest, montane, steppe, and desert forms. They are 
of relatively moderate significance as fur animals or as exhibits for zoological 
parks. Due to persecution the population of some species has fallen shaфly 
and these are now on the verge of extinction. One species (tiger) has suffered 
a particularly dramatic reduction in population and range and is placed under 
conservation; two other species (cheetah and caracal) are under protection. 

63 Key for Identification of Species of the Cat Family, Felidae^^ 

Identification by External Characters 

1 ( 2). Sharp, vivid, dark (black) band extends from inner comer of eye 
along nose toward upper lip on a light field. General color very 
light, yellowish with small, black, dense, circular spots not forming 

rings. Claws project significantly from fur covering paw 

cheetah [gepard]* Acinonyx jubatus (p. 702). 

2(1). Well-developed black band extending from inner corner of eye 
to upper lip absent. General color different. Claws do not project 
from fur covering paw. 

3 ( 4). Vivid vertical (transverse) black stripes extend through bright 

reddish background color on back and flanks. Very similar pattern 
of stripes occurs on head. Size very large (body length of adults 
exceeds 160 cm) tiger [tigr] Panthera tigris (p. 95). 

4 ( 3). Color different. Size smaller. 

5 ( 8). Groups of black spots arranged in form of rings dispersed 

throughout light background together with individual dense spots 

^-Currently labeled Felis silvestris L. forma catiis. See the section "Description" under 
steppe cat (F. s. caudata) for the origin of domestic cats. 

^■'Since the Hon is no longer found in the Soviet Union, it is not included in the key. 
"Transliteration of Russian common names in brackets. 


of moderate size. Size fairly large (body length of adults exceeds 
90 cm, often over 100 cm, and even 120 cm). 

6 ( 7). Main color of trunk light yellow or reddish yellow. Tail length 

less than length of head and body. Spots shaфly outlined and vivid 
black .... leopard [bars, "leopard"] Panthera pardus (p. 203). 

7 ( 6). Main color not yellow but dirty white or very light gray. Tail length 

roughly equal to length of head and body. Spots, at least in winter 

coat, not vivid and not sharply profiled 

snow leopard, ounce [snezhnyi bars, irbis] Uncia uncia (p. 276). 

8 ( 5). Color different. If spots present not arranged in form of large rings. 

9 (10). Tail length, without terminal hair, less than that of hind foot or 

equal to it; terminal one-third or one-fourth of tail black 

lynx [rys'] Feiis lynx (p. 524). 

10 ( 9). Tail length, without terminal hair, more than length of hind foot. 

Tail end not black or black for less than one-third or one-fourth 
of tail length. 

1 1 (12). Two distinct light (white) bands, usually bordered by black, extend 

from inner comers of eyes along forehead. Back of ears with well- 
defined white field (spot). Ocherous or reddish spots or transverse 

band visible on throat and lower surface of neck 

. . . leopard cat, Amur cat [amurskii kot] F. euptilui'a (p. 328). 

12 (11). White bands extending from inner comers of eyes along forehead 

absent; rear of ear without white spot; reddish or ocherous spots 
and transverse bands on throat and lower surface of neck absent. 

13 (16). Spots distinctly visible. 

14 (15). Generaltoneofbackandflanksbrownish-gray. Along back, double 

line of merged spots visible and similar vertical bands present 
on sides in addition to individual spots. Throughout its length 
tail uniformly covered with long hair (not thinned toward the end, 
terminal hair long) and appears trimmed. Tail with several (3 to 

10, usually 5 to 7) black transverse bands (Fig. 20)* forest 

wild cat [Evropeiskaya lesnaya koshka], Fells silvestris, group 

silvestris (p. 402). 

64 15 (14). General tone light dirty gray or pale gray. Back and flanks covered 

with innumerable distinct spots. Double black band along back 

and transverse stripes on flanks not well defined, or ver)^ faint. 

Tail tapers toward end (Fig. 20) steppe 

wild cat [stepnoi kot], Felis silvestris, group libyca (p. 441). 

16 (13). Body monochromatic, without distinct spots, or with several dark, 

narrow, transverse bands running across back. Dark marks (spots 

^'Erroneously given as 19 in Russian original — Sci. Ed. 


64 Fig. 20. Form and pattern of tail of European (and Caucasian) forest wildcat, Felis 

{Felis) silvestris, group silvestris and steppe wildcat, F. (F.) silvestris, group libyca. 
Collection of the Zoological Museum, Moscow University. Sketch by N.N. 


or bands) may occur on legs; if some individual spots occur on 

body, they are blurred, indistinct, and small. 

Back of ears black; tuft of long (usually over 20 mm) hair at tip 

of ears caracal [karakal], Felis caracal (p. 498). 

Back of ears not black; tuft on ears absent, or hair on tip of ears 

shorter (less than 20 mm). 

Lower surface of paw with long, dense, black or blackish-brown 

hair covering sole and digital pads 

sand cat [barkhanaya koshka] Felis margarita (p. 636). 

.*Lower surface of paw devoid of long, dense, black hair and sole 
and digital pads hairless. 

,*Tail relatively short and does not, or barely reaches calcaneal 
joint. Long hair on cheeks (whiskers) absent. Hair at tip of ears 
sometimes slightly elongated. Ears large, with pointed tips, and 

project beyond fur in winter 

junglecat [kamyshovyikot]FeIischaus(p. 356). 

22 (21).*Tail relatively long, end extending significantly beyond calcaneal 











*Misnumbered in Russian original — General Editor. 


joint. Long hair (whiskers) present on cheeks. Hair at tip of ear 
not elongated. Back of ear with blurred, narrow, transverse bands. 
Ears short, rounded above, and project very little from dense rich 
coat in winter. . . . manul cat [manul]. Fells manul (p. 655) (V.H.) 

Identification by Skull Characters ^"^ 

1 ( 8). Size large. Length of upper camassial tooth more than 21 mm. 

2 ( 3). Length of upper camassial tooth 29 mm or more. Condylobasal 

length 250 mm or more tiger, Panthera tigris (p. 103). 

65 3 ( 2). Length of upper camassial tooth less than 29 mm.^-** 
Condylobasal length not more than 240 mm. 
4 ( 5). Maximum width of inteфterygoid vacuity more than least distance 
between tympanic bullae. Inner cusp of upper camassial tooth 
not developed or barely visible and devoid of peak (tubercle) (Fig. 
21). Distinct small cusp present in front of main cusp of third 

premolar (before camassial) (Fig. 22) 

cheetah, Acinonyx* jubatus (p. 702). 

Fig. 21. Upper camassial tooth o( cheetah, Acinonyx jubatus (left) and snow leopard, 

Uncia uncia. Nos. S 47284 and S 49240, collection of Zoological Museum, Moscow 

University. Sketch by N.N. Kondakov. 

^""On examination of material some of the "traditional" features used for the 
identification of this genus were found to be inconsistent; namely, form of infraorbital foramen 
was sometimes, and in some species often, double; most features of relative disposition of 
posterior end of nasals and frontal processes of upper jaw; condylobasal length; and other 
major features, viz., position of anterodorsal end of jugal in relation to lacrimal duct. In 
the key given here an attempt has been made to include some new features together with 
some fairly common ones. 

^^Applies to permanent and not deciduous tooth. Reliable identification of juveniles, 
especially from deciduous dentition, is possible only in museums with significant comparative 

*Misnumbered/misspelled in Russian original — Sci. Ed. 


Fig. 22. Third and fourth (carnassial) premolars of snow leopard, Uncia uncia 

(top) and cheetah, Acinonyx jubatiis. Nos. S 47284 and S 49240, collection of 

Zoological Museum, Moscow University. Sketch by N.N. Kondakov. 

5 ( 4). Maximum width of inteфterygoid vacuity less than least distance 

between tympanic bullae. Inner cusp of upper carnassial tooth 
well developed and with prominent peak (tubercle) (Fig. 21). 
Distinct cusp in front of main cusp of third premolar (before 
carnassial) absent (Fig. 22). 

6 ( 7). End of paroccipital process extended, overhanging rear of tympanic 

bulla in the form of very faint pointed hook (Fig. 23*). Antero- 
lateral (ectotympanic) chamber of tympanic bulla well developed 
(Fig. 23). Anterior wall of swollen portion of bulla almost at level 
of posterior wall of glenoid fossa (postglenoid process; Fig. 23) 
snow leopard, Uncia uncia (p. 284). 

7 ( 6). Paroccipital process short, thick, and blunt; adjoins posterior wall 

of tympanic bulla completely; no long, thin, overhanging end in 
form of a hook (Fig. 23). Almost all of bulla consists of postero- 
medial (entotympanic) chamber; anterolateral (ectotympanic) 
portion very small (Fig. 23). Anterior wall of swollen portion of 
bulla separated from posterior wall of glenoid fossa (postglenoid 
process) by a distance roughly equal to width of bulla (Fig. 23) 
leopard, Panthera pardus (p. 208). 

8 ( 1). Size small. Length of upper carnassial tooth less than 21 mm. 
66 9 (10). Length of upper camassial tooth 17 mm or more.^^ Second upper 

*Misnumbered in Russian original — Sci. Ed. 

^^In rare cases, when length of upper camassial tooth is about 17 mm (± 0.1 to 
0.2 mm), the character should be compared with those given in parentheses in 15 (16). 


\^ . 



66 Fig. 23. Tympanic bulla and paroccipital process in snow leopard, Uncia iincia 

(left) and leopard, Panthera pardus. Upper row — ventral view; lower row — 
lateral. Nos. S 35132 (snow leopard) and S 51945, collection of the Zoological 
Museum, Moscow University. Sketch by N.N. Kondakov. 

premolar (second tooth before camassial) absent (three teeth occur 
in upper jaw behind canine). ^^ lynx, Felis lynx (p. 532). 

10 ( 9). Length of upper camassial tooth less than 17 mm. Second premolar 

in upper jaw present or absent (three or four teeth occur beyond 

1 1 (14). Ectotympanic chamber of tympanic bulla very large; bullae greatly 

swollen and very large; its anterior point at level of posterior wall 

^^As a rare exception, very small second premolars may be present but usually only 
on one side. 





Fig. 24. Tympanic bullae of manul, Felis mamil (left) and sand cat, F. margarita 
(center) with highly developed anterolateral (ectotympanic) chamber. Right, for 
comparison, bulla of steppe wildcat F. silvestris, group libyca, with a verj' small 
anterior chamber. Nos. S 40113, S 64765 and S 42325, collection of the Zoological 
Museum, Moscow University. Sketch by N.N. Kondakov. 

of glenoid fossa (postglenoid process) or even slightly shifted 
forward (Fig. 24). 

12 (13). Inner cusp of upper carnassial tooth well developed and with 

distinct small peak (tubercle; Fig. 25). Maximum diameter of 
external auditory meatus roughly equal to length of upper 
carnassial tooth. Second premolar present in upper jaw (four teeth 
behind canine). ^^ sand cat, Felis margarita (p. 642). 

13 (12). Inner cusp of upper carnassial tooth not developed and peaks 

(tubercles) not evident (Fig. 25). Maximum diameter of external 
auditory meatus less than length of upper camassial tooth. Second 

premolar on upper jaw (three teeth behind canine) . 

manul cat, Felis manul (p. 671). 

14 (11). Anterolateral (ectotympanic) chamber of tympanic bulla small 

and almost all of bulla formed by posteromedial (entotympanic) 
chamber. Anterior point of tympanic bulla does not reach level 
of posterior wall of glenoid fossa (postglenoid process). 

15 (16). Distance between tips of supraorbital processes only twice the 

interorbital width. Distance between supraorbital process of frontal 

Fig. 25. Upper carnassial tooth of manul, Felis manul (left) and sand cat, F. 

margarita. Nos. S 40113 and S 67765, collection of the Zoological Museum, 

Moscow University. Sketch by N.N. Kondakov. 

'^Sometimes this tooth, especially in old animals, is lost from one or both sides but 
fairly distinct traces of its alveolus invariably remain. 


67 Fig. 26. Shape of nasal process of premaxilla and its relation to nasal process 
of frontal in caracal, Felis caracal (left) and Caucasian wildcat, F. s. caucasica. 
Nos. S 40195 and S 17640, collection of the Zoological Museum, Moscow 

Univereity. Sketch by N.N. Kondakov. 

process of premaxilla narrow and long, and tapers gradually 
upward; tip elongated and sharp; extends closer to forward 
projection of frontal; distance between them usually not more than 
length of third upper premolar (Fig. 26*). Second upper premolar 
usually absent. (A distinct bony crest runs to orbit along outer 
side of pterygoid bone. At its posteriormost end it forms a broad 
plate in the form of a triangle or fork. Length of tympanic bulla 
measured from canal [foramen] of carotid artery to the point of 
its union with the lateral occipital process is equal to distance 

between ends of upper canines or slightly more.)^^ 

caracal, Felis caracal (p. 503). 

16 (15). Distance between ends of supraorbital processes more than two 
times interorbital width. Orbit closed from rear or distance between 
supraorbital process of frontal and postorbital process of jugal 
less than length of latter. Nasal process of premaxillae relatively 
short, broad along much of its length, and constricted upward in 
form of a relatively blunt wedge. Distance between its tip and tip of 

68 forward projection of frontal more than length of third upper 
premolar (Fig. 26). Second upper premolar usually well developed. 
Posterior margin of bony palate outside interpterygoid vacuity 
notched, steep, and deep. 

*Misnumbered in Russian original — Sci. Ed. 
^^See footnote No. 16 above. 


Fig. 27. Shape of nasals in jungle cat, Fclis chaus (left) and steppe cat, Felis 

silvestris, group llhyca. Nos. S 3603 and S 42325, collection of the Zoological 

Museum, Moscow University. Sketch by N.N. Kondakov. 

17 (18). Length of nasal, measured from posterior end to anterior projection, 
more than distance from posterior margin of anterior palatine 
foramen to posterior margin of bony palate at a corresponding 
point in interpterygoid vacuhy, or exceptionally, equal to it. 
Posterior ends of nasals usually have an extended, pointed shape 
(Fig. 27) jungle cat, Felis chaus (p. 362*). 

Fig. 28. Position of posterior margin of bony palate in interptery-goid vacuity in 

Amur cat, Felis euptilura (left) and European (Caucasian) wildcat, Felis silvestris. 

Nos. S 14261 and S 75098, collection of Zoological Museum, Moscow University. 

Sketch by N.N. Kondakov. 

*Misnumbered in Russian original — Sci. Ed. 


18 (17). Length of nasal, measured from posterior end to anterior projection, 

less than distance from posterior margin of anterior palatine 
foramen to posterior margin of bony palate at a corresponding 
point in interpterygoid vacuity, or exceptionally equal to it. 
Posterior ends of nasals have a tapered, only slightly extended 
shape (Fig. 27). 

19 (20). Bony palate long; its posterior margin in region of inteфterygoid 

vacuity lies at level of posterior surface of molars at a distance 
roughly equal to length of third premolar or at a distance roughly 
equal to one- half or more of width of inteфterygoid vacuity (Fig. 
28). Interpterygoid vacuity of uniform width and narrow;* its inner 
width in anterior part not more than 10 mm and width along outer 

walls at level of anterior margin less than 15 mm 

Amur cat, Felis euptilura (p. 335). 

20 (21). Bony palate short; in region of pterygoid [sic] vacuity its posterior 

margin lies at level of posterior surface of molars, or only slightly 
beyond this line, at a distance much less than length of third 
premolar and less than one-half of width of inteфterygoid vacuity 
69 (Fig. 28). Inteфterygoid vacuity somewhat broader anteriorly than 

posteriorly; broad,* its inner width in anterior part exceeds 10 
mm and width along outer walls, at level of anterior margin, 
exceeds 15 mm. 

21 (22). Maximum diameter of external auditory meatus more than length 

of third upper premolar (lies before carnassial tooth) (Fig. 29) . . . 
steppe wildcat, Felis silvestris, group libyca (p. 450). 

22 (21). Maximum diameter of external auditory meatus less than length 

of third upper premolar or equal to it (Fig. 29) 

forest wildcat, Felis silvestris, group silvestris (p. 407). (V.H.) 

Fig. 29. Tympanic bulla and external auditory meatus of steppe wildcat, Felis 

silvestris, group libyca (left) and European (Caucasian) forest wildcat, F. s., group 

silvestris. Nos. S 42325 and S 17640, collection of the Zoological Museum, Moscow 

University. Sketch by N.N. Kondakov. 

"Appears inconsistent with Fig. 28, but not Figs. 158 and 183 — Sci. Ed. 


Genus of Large Cats 

Genus Panthera Oken, 1816 

1816. Panthera?^ Oken. Lehrb. Naturgesch., 3, 2, 1052. Felis pardus 

1816. Tigris, Oken. Ibid., 3, 2, 1016. Felis tigris Linnaeus. 
1816. Leo. Oken. Ibid., 3, 2, 1070. Felis leo Linnaeus. 
1829. Leo. Brehm. his (Oken), p. 637. Felis leo Linnaeus. 
1843. Tigris. Gray. List Mamm. Brit. Mus., p. 40. Felis tigris Linnaeus. 
1868. Pardus. Fitzinger. Sitz.-Ber. K. Ak. Wiss. Wien, 58, 1, 459. Felis 
pardus Linnaeus. (V.H.) 
Large to very large animals; largest in the family. 

70 Fig. 30. Body contour of big cat of genus Panthera and small cat of genus Felis. 

Elevation of sacral region in small cats is characteristic. Depicted — a lioness and 
a domestic cat. Sketch by N.N. Kondakov. 

^*^his name does not fully satisfy the rules of nomenclature but is extensively used. 
It should be retained as a nomen conservandum. 


General appearance wholly typical for all cats. Legs in most not long, 
but thick; trunk extended, sometimes highly so. Sacral region not high and 
back not raised toward rear; either straight or height at shoulder (withers) 
more than at sacrum or back slopes gently toward rump (Fig. 30). Tail long, 
not less than one-half of body length, or slightly more. Head moderate in 
size or large, somewhat extended with a relatively elongated facial region, 
and fairly straight or faintly arcuate dorsal profile. Ears relatively small, 
short with a blunt rounded tip, and wide-set. In male lions and animals 
in winter coat ears project slightly out of coat. Long hair (tufts) on top 
of ear absent. Eyes relatively small and pupil round. 

Pelage in most species uniform throughout body, and tail uniformly 
fiirred throughout its length. Sometimes hair on cheeks elongated (whiskers); 
in one species (lion) well-developed mane covers anterior portion of body 
and tail with long tuft at end. Legs strong with broad, powerful paws, 
especially forepaws; claws large, very shaф, sharply bent, and fully retractile. 
At end of all digits, dermal thickening occurs above and below claws; this 
thickening completely conceals retracted claw. Interdigital membranes large 
and reach end of digital pads (Fig. 278). In most species hair short and not 
dense, becoming dense and relatively long only in extreme northern forms 
of some species in winter. Color monochromatic, with black transverse 
stripes, or with black spots dispersed against light-colored background as 
single spots or groups forming rosettes. In most species large white spot 
occurs on back of ear. 

Teats, two or three pairs. 

Skull large and heavy, with massive bones, thick and powerful 
zygomatic arches, and powerfully developed tuberosities and crests. Sagittal 
crest particularly long, very high and projects notably backward; lambdoidal 
crest well developed. Dorsal profile fairly straight or only faintly and 
uniformly convex; interorbital region raised relatively weakly; line of profile 
slopes gently back from it. On the whole skull appears flattened. Basicranial 
axis (floor of cranium) almost horizontal and does not form a distinct angle 
with basifacial region. Depression (inflexion) at point of transition of nasal 
into interorbital region on upper part of skull absent. 

Brain case of relatively moderate size, very small in some, and elongated 
as though greatly pressed by masticatory muscle attached to sagittal crest. 
Postorbital constriction marked; sometimes its width is equal or even less 
than width of interorbital constriction. Facial region of skull very powerful 
and broad (in region of zygomatic arches and in region of canines) and also 
elongated. Measuring along a transverse line passing through the supraorbital 
71 processes, facial region larger, or not less than, braincase as a whole, or 
without portion of sagittal crest extending beyond occiput. 

There is fairiy large separation, roughly equal to width of tympanic bulla. 


between postglenoid process and bulla. Tympanic bullae relatively small, 
not bulging, flattened from ventral side, and thick-walled (small m leopard, 
but stands out); inner chamber (entotympanic) significantly larger than outer 
(ectotympanic or tympanum proper), and septum between them lies close 
to outer ear passage. In hyoid apparatus (Fig. 31) third upper element 
(ephihyal) not ossified and forms a hinge ligament which permits freer 
movement of the larynx (see below). Deep depressions absent or barely 
perceptible on basioccipital portion of occipital bone close to tympanic bulla 
in front of jugular foramen. 

Dentition complete; second premolar (first in row) usually present on 
upper jaw. Teeth very powerful; canines relatively short but strong with 
a broad base. 

Sexual dimorphism in most species manifested in large size of males 
and in part more intense development of crests, etc. on skull; age-related 
color dimoфhism not evident. In one species (lion) male differs 8Ьаф1у 
in appearance in growth of mane on anterior part of trunk and elongated 
hair at some places on chest and abdomen; adults monochromatic but color 
of newborn and young spotted. Seasonal dimorphism in color absent; winter 
coat denser and with long hair in northern or hill subspecies of some 

These predators feed mainly on big mammals (ungulates). The prey 
is usually bigger than the predator, sometimes by several times. Hunting is 

70 Fig. 31 . Hyoid apparatus (sketch) in genus of big cats, Panthera Oken (left), and 

genus of small cats, Felts L. and cheetah, Лсшонуд:. Sketch by N.N. Kondakov 
from H. Hemmer. 


done by concealment and ambush on pathways, at water holes, and so on. 
They are noctumal animals, and with one exception solitary. The male does 
not participate in the feeding of the young. One species (lion) lives and 
hunts in small groups, representing a family unit, as in the case of wolves 
(see Volume II, Part 1). They are mostly inhabitants of plains and montane 
forests, sometimes open forestless mountains, and reed thickets; one species 
lives in open steppes (savannas) and semideserts. 

As a result of the specialized nature of the hyoid apparatus (see above), 
the larynx and vocal chords are highly mobile and the voice a loud coarse 
72 roar ("roaring cats"; big cats are incapable of "meowing"). They do not 
express contentment and well-being by "purring", or "purr" only on 
exhaling. They mainly eat while resting on the abdomen, keeping the forearm 
and ulnar region in contact with the ground, and tear the prey to pieces 
by jerking the head upward (Fig. 32). 

The range of the genus (reconstructed) (Fig. 33) includes Africa, extreme 
southeastern Europe, Asia (except the northern part), and South, Central 
and southernmost North America. In South America the range covers all 
of the northern and partly the central part of the mainland; southward it 
includes Peru, Bolivia, Paraguay, and southern Brazil roughly up to Sao 
Paulo, the province of San Luis, and even La Pampa in Argentina (slightly 
north of 40° NL). Northward the range encompasses all of Central America, 

<1Ш i/ 

Fig. 32. Usual posture of a big cat of genus Panthera while eating (forearm region horizontal 
to substratum) and small cat of genus Felis (ulna raised high). Sketch by N.N. Kondakov. 

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rougMy up to the latitude of Veracruz in southern Mexico; and farther, 
bypassing on the east and west the Mexican highlands, extends along the 
lowlands of the Pacific Ocean and the Caribbean Sea, and extends in the 
form of two long extensions into southern Texas on one side and California, 
Arizona, and New Mexico on the other. The California Peninsula [i.e., Baja 
California] proper and all islands of the western hemisphere fall outside 
the range. 

The range encompasses all of Africa and the southern parts of the Balkan 
Peninsula in Europe. In Asia the southem boundary encompasses the Arabian 
Peninsula, runs along the southern edge of the mainland, and covers Sri 
Lanka, Malacca, Sumatra, Java, Bali, and bCangean Island (north of Bali). 
In the east the boundary similarly runs along the coast of the mainland but 
species of the genus are not found in Hainan,^^ Taiwan, or the islands of 
Japan and Shanter Islands. In Sakhalin only stray animals have been reported 
(tiger). In northern Asia the range includes the Amur, Trans-Baikal, Altai, 
and Kashgar, eastern and southem Kazakhstan, and Middle Asia, Iran, the 
Caucasus, and Asia Minor (for details see distribution of tiger and leopard). 
Over the historic period the range, at places, has shrunk due to extinction 
(Europe, Middle Asia, Siberia, parts of Africa and southem Asia, and others) 
and continues to shrink even more rapidly elsewhere. 

Species of this genus are among the best studied in the family, especially 
their moфhology and ethology.22 Differences of opinion among syste- 
matists relate only to the relative progressiveness of individual species and 
their division into subgenera (see below). 

As already mentioned above, there is justification to regard big cats 
as less specialized among genera of the family. In any case, compared with 
other groups of the family (different groups of species in genus Felis) in 
which feline type specialization has reached utmost development and 
expression, members of this genus represent a less specialized, more 
generalized "predatory" type in skull structure. It is possible that some 
features of species of genus Panthera are secondary (allometric). 

Big cats appeared in the Early Pliocene of Eurasia. In the Early 
Pleistocene of Eurasia they were represented by special species (P. 
arvemensis and P. cristata), sometimes considered the ancestral forms of 

^^V. Mazak (1965, p. 33) has indicated (arrow) on the distribution map of tiger he 
presents, a sighting on Hainan. This is clearly erroneous. Mazak evidently had in mind old 
infoiTnation about the intrusion of tiger from the mainland onto Amoy Island (5 miles from 
the mainland) cited by Allen (1938). There is no positive reference whatsoever in literature 
(including also Mazak, 1965) to Hainan. 

^"Literature devoted to this group of cats is extensive. The basic works are those of 
Severtsov, 1858; Pocock. 1917; and others; Haltenorth, 1936 and 1937; and others; Weigel, 
1961; and others; and the bibliographic compilation of Hemmer, 1966. 


lion and tiger. The cave lion or "tiger-lion" (P. spelea), extensively 
distributed throughout the whole of the Pleistocene period, is an independent 
species with which present-day lion and tiger have no direct genetic 
association. In the New World the genus is known only in the Pleistocene 
period (giant jaguar of North America, P. atrox). 

There are four present-day species in the genus: tiger, P. dgris Linnaeus, 
1758; lion, P. leo Linnaeus, 1758; leopard, P.pardus Linnaeus, 1758; and 
jaguar, P. onca Linnaeus, 1758. These constitute 11.5% of the species of 
the family. 

All these forms constitute a group which is moфhologically and 
phylogenetically very close, but views on the affinities between the various 
74 species are quite diverse. Some (Pocock, 1929, 1930, and 1941) do not 
divide the genus into subgenera, or assuming a large number of species 
in the genus (Haltenorth, 1936 to 1937) place the foregoing species in a 
single subgenus; other authors (Hemmer, 1966) classify them into two 
subgenera by placing tiger in a subgenus, treating it as the most primitive 
form, and the remainder in a special subgenus (Ellerman and Morrison- 
Scott, 1956 and 1966; see list of synonyms). Most taxonomists consider 
the lion the most primitive form of the genus and see in the tiger the species 
closest to lion. It has, however, been suggested (Hemmer, 1960), based 
on the sum total of moфhological and ethological features, that the tiger 
should be regarded as the most primitive and lion the most specialized form 
of the genus. In the present work a purely moфhological series — lion- 
tiger-leopard — has been adopted without classifying them into subgenera. 
The maximum number of species (three) occur in the Old World, one in 
Africa and Asia (leopard), one in Africa, Asia, and Europe (lion), and one 
purely Asian (tiger). There is only one species (jaguar) in the New World. 

Panthera are very valuable game animals and their fur highly prized. 
They regulate ungulate populations and at places destroy agricultural pests 
(tiger, wild boar, African leopard, baboon). In some areas they also destroy 
cattle and pose a danger even to man. 

Russian fauna includes three species: lion, P. leo Linnaeus, 1758; tiger, 
P. dgris Linnaeus, 1758; and leopard, P. pardiis Linnaeus, 1758. These 
comprise 75% of species of the genus, about 11.5% of species of the family, 
and about 1.0% of species of Russian fauna. Lion had already become totally 
extinct in the tenth century. 

The range covered the southern montane and forest region, at places 
semidesert and desert parts of the country, and has shrunk markedly in the 
last decade. These are forest and montane forms, and inhabitants of reed 
thickets. In view of their small numbers, their economic importance is 
negligible and rarely adverse. One species (tiger) is totally protected; like 
the leopard, it is of much interest to zoological gardens. (V.H.) 


Panthera leo Linnaeus, 1758 

1758. Fells leo Linnaeus. Syst. Nat., 10th ed., vol. 1, p. 41. Constantine, 

1826. Felis leoperslcus. Meyer. Diss, inaug. anatomicomedica de Genera 

Felium, p. 6. Persia.^ (V-H.) 


Animals of large size (one of the two largest species of the family). 

General appearance typical of big cats. Body greatly extended, tail length 
about one-half of body length, and height at sacmm not more than height 
at shoulders. Head relatively large and heavy, and markedly extended (with 
a large facial region). Legs strong with broad and powerful paws. Tail with 
tuft of long (black) hair at end (Fig. 34). 

Sexual dimorphism very pronounced, much more distinct than in any 
other species of the family, and very characteristic. Coat of females even, 
short, fairly stiff, and uniformly dense sandy- or brownish-yellow color 
without pattern. 

The difference in color of flanks and abdomen, and outer and inner 
surfaces of legs, the latter being somewhat lighter, is insignificant and devoid 
of sharp delineation. Black spot occurs on back of ear. Hair on flanks and 
upper surface of neck turns up and forward, and elsewhere on the body 
76 downward. Lower surface of tmnk and inner surface of legs usually with 
blurred pale spots, darker and more rust-colored than main background, 
but faint. Frequently such spots are also visible on flanks and even form 
very faint vertical rows. Black tuft at end of tail somewhat smaller in female 
than in male. Rarely, mdimentary mane visible in very old females, con- 
sisting of very short and sparse hair covering a small area on the head 
and neck. 

Males and females are similar in color but the former have a large mane 
of dense hair which, in old animals, reaches 40 cm in length. The mane, 
although forming a single unit, can be divided into several parts: one segment 
(chest mane) includes rear of head (commencing from ears), sides of neck 
both above and below, longer hair on sides of head (whiskers), and a large 
mass on chest and between forelimbs. Elongated hair (abdominal mane) 
extends in the form of a band along the abdomen (sometimes along its entire 

■^Additional synonyms iiave been given by R. Pocociv (1930 and 1939). 






length). Another broad band of long hair covers the underside of the flanks 
from the rear of the axilla up to the groin region (lateral mane). Hair on 
the rear of the forelegs from the ulna downward is likewise elongated. It 
is longest on the ulna and often forms tufts; below the ulna it is shorter 
and finally disappears. From the neck and nape long hair extends backward, 
covering the back in the region between the shoulders (shoulder mane) or 
even slightly farther back. It may cover the upper part of the shoulder region 
or form a cape along the rear of the scapular region. This cape may extend 
downward so far that it merges with the abdominal and lateral manes 
(see Fig. 39). 

Mane growth commences at the age of six months. Gradually increas- 
ing in density, size, and luxuriance, the mane is fully grown by the 
third year but attains maximum growth and luxuriance in old animals. 
The mane consists of yellow-colored hair, corresponding to the general 
color of the trunk, with an admixture of more or fewer black hair, and 
some of lighter color — white, or very light yellow hair. The latter are 
more common in the frontal (facial) portion of the male. A large part 
of the mane, mainly the rear portion, is black and darkens even more 
with age. 

The general luxuriance, density, and color of the mane, as well as the 
extent of its growth (form and area covered) exhibit individual variability. 
Geographic variability is also significant. The most luxuriant and extensive 
manes (tresses on flanks and abdomen) occurred in extinct lions of the Cape 
region (P. /. melanochaitus) and northwestern Africa (Atlas; P. I. led). The 
mane is less developed in other forms, and in some covers only the head, 
neck, and front of the chest.^'^ 

Faint spots are sometimes seen on males, but rarely so; the spots are 
vague and do not cover large areas of the body. They are seen only on 
the abdomen and paws and never on the flanks; in rare cases traces of them 
might be discerned on the latter. The final feature of sexual dimorphism 
in lion, as in all cats, is manifested in the smaller size and much lighter 
build of females. 

Newborns are, as a rule, covered in distinctly circular dark spots, which 
tend to form vertical rows; in some animals these spots even fuse into distinct 
vertical stripes. Sometimes newborns are monochromatic and have only 
faint dark spots on the underside. Usually this skin pattern has gradually 
disappeared by the age of about six months; more rarely it is preserved 
for two to three years or even longer. Females have been known to preserve 
abdominal skin spots throughout their life. 

^"^A particularly luxuriant mane is grown by caged lions. Tlie mane of wild animals 
is often damaged by their movement through underbrush and bushes. 


The skull is very similar to that of tiger (see below) but usually some- 
what more depressed and flattened in the frontal region, with a slightly shorter 
postorbital region. Hence the facial part, compared with the cranial, appears 
longer and more massive than in tiger. The nasal openings are usually broader 
77 but the nasals quite short, not projecting beyond the rear of the maxillae 
(Fig. 35). The inner cusp of the upper carnassial tooth is very small. The 
lower profile of the lower jaw bulges and hence when placed on a flat surface 
the jaw rocks; contrarily, the lower jaw of a tiger is concave and does not 
rock. These features differentiate the skull of these two species, but the 
individual variability of tiger skulls is so great that only the structure of 
the lower jaw constitutes a reliable feature for positive differentiation^'' 
(Fig. 36). 

In general body dimensions and skull the lion is similar to the tiger. 
The weight of an adult male lion is usually about 175 kg and of a lioness 
about 125 kg. However, much heavier animals are not rare. Large South 
African male lions weigh 180 to 226 kg and females 127 to 140 kg. Body 
length of males about 170 to 180 cm and tail length about 70 to 85 cm. 
Maximum skull length in males 321 to 401 mm, females 292 to 333 mm; 
condylobasal length in males 309 to 348 mm, females 263 to 291 mm; 
and zygomatic width in males 220 to 256 mm, females 188 to 212 mm 
(data from Roberts, 1959). (V.H.) 

Systematic Position 

There is no doubt that the lion is closer to the tiger than to any other cat. 

Fig. 35. Relarionship between nasal and frontal bones in lion (above) and tiger. 
Sketch by N.N. Kondakov. 

•^^Pocock (1939) thought that this feature was "absolutely constant". However, 
Haltenorth (1936) established that in roughly 10% of lions (11 skulls out of 113), the lower 
profile of the lower jaw was either straight or even concave. 


Fig. 36. Lower jaw and line of its ventral profile in lion (upper line) and tiger. 
Sketch by N.N. Kondakov. 

Their identical skull structure bears this out. Both species are evidently 
the least specialized forms of the genus and lion is usually placed at the 
head of the series (Haltenorth, 1936-1937 and 1957), although some authors 
begin the series of big cats with tiger (Pocock, 1939; and others). Recently, 
a classification of the genus has been proposed according to which tiger 
is separated in one subgenus, at the beginning of the series, while jaguar, 
leopard, and lion are placed in another subgenus with lion at the end of 
the series, i.e., the two species are widely separated and represent terminal 
species of the genus (Hemmer, 1966). In building up this scheme, ecological, 
ethological, and genetic data have been used, but it is not well substantiated 
moфhologically and is hardly acceptable at present. (V.H.) 

Geographic Distribution 

The range (reconstructed) encompasses the open, often desert, expanses 
of Africa, the Middle East, southeast Europe, and northern India. The range 
78 is greatly reduced at present. 

Geographic Range in the Soviet Unioti 

The range (reconstructed) in the Soviet Union is very small and covers 
only the eastern part of the Trans -Caucasus (Fig. 37). 

Lions are known from the Trans-Caucasus from the Holocene and 
became extinct only in the tenth century. In the Trans-Caucasus this cat was 
not incidental (for example, rare migrations) but rather an important member 
of the fauna. This is supported by the dispersal of the animal in the past 
into regions adjoining Trans-Caucasus in the south. These regions formed 
a single faunal complex, with the Trans -Caucasus representing the northern 


Fig. 37. Probable maximum area of distribution of lion, Panthera leo L. in the 
Caucasus in the Holocene (from N.K. Vereshchagin, 1959, with modifications). 

V.G. Heptner. 

border. Commencing evidently in the first centuries of our era, the inhabited 
area and numbers of the animal steadily decreased in the Trans-Caucasus. 
However, even in the early Middle Ages lions were evidently fairly common 
and hunted by shirvanshakhs [native people of Trans-Caucasus]. The total 
disappearance of the animal from the Soviet Union occurred in the tenth 

To establish a reasonably precise distribution for the lion is now very 
difficult. Information in the literature and data available for the Middle Ages 
and folklore traditions are little indeed. During the period of its greatest 
extent the range covered all the plains and the foothills of the eastern Trans- 
Caucasus and extended westward almost to Tbilisi. In the north, bypassing 
the eastern part of the Caucasus, it formed a large extension from the 

"^^Information about the occurrence of lion, together with tiger, in the mid-1600's in 
western Georgia (Mingreliya and Imeretiya) is erroneous (Vereshchagin, 1959). 


Apsheron Peninsula to the mouth of the Samur [river]. Extending then to 
Araks, the boundary turned east in the form of a narrow projection to Yerevan 
and a little farther. Its northern boundary then extended in the west into 
79 From the general nature of the range of lion in the Middle East (see 

below), intrusions of the animal in the past into the extreme south of 
Middle Asia, primarily in southern Turkmenia and probably along upper 
Amu-Darya, cannot be excluded. This is supported by some geographic 
names involving the term "shir", i.e., lion, for example Shirabad, a town 
on the Shirabad-Darya, a tributary of the Amu-Darya, or the Shir-Tepe 
region on the upper Tedzhen, and others. In fact, there is no direct 
historic proof of the existence of lion in Middle Asia but it is possible 
that a study of the source books of the Middle Ages could provide such 
information. The range of the lion in Persia in antiquity stood quite close 
to the boundaries of Turkmenia, and the cat evidently prowled even in 
Afghanistan. Habitat conditions of these regions even in the early twentieth 
century were favorable for the survival of big predators (tiger, cheetah, 
wolf, hyaena, and leopard). In the past, and more so in the very remote 
past, extensive tugais, pistachio savannas, rushes, etc., together with an 
abundance of kulan, goitered gazelle, wild boar, mountain sheep, and 
deer constituted an environment which was even more luxurious for the 
survival of lion. 

Various views have been expressed about the occurrence of lion, or 
at least its appearance, in the southern European parts of the USSR. These 
views are based on the inteфretation of the old Russian "lyuti zver' " (fierce 
animal). The well-known citation, always referred to, are the words of the 
great Count of Kiev, Vladimir Vsevolodovich Monomakh (1053 to 1125) 
in his work, Poucheniya Detyam (1117). In describing a hunt in the years 
of his rule in Turov and Chernigov (1073 to 1094), he wrote that a fierce 
animal sprang toward his thighs, injuring both him and his horse. The 
traditional interpretation, suggesting either a wolf or lynx as this "fierce 
animal", is evidently unacceptable; these animals could hardly spring at 
the thighs of a rider, let alone throw a horse. At present some think this 
"fierce animal" was a leopard while others favor lion. The latter 
interpretation (Charlemagne, 1964^^, and others) is quite widely accepted. 
However, there is undoubtedly a good basis for seeing in this "fierce animal" 
a tiger (see chapter on tiger below; Heptner, 1969). Thus the appearance 

^^The distribution and liistory of lion in Trans-Caucasus are mainly based on data 
from N.K. Vereshchagin (1959). 

^^liis work includes a list of the main publications on this subject; also see V. 
Mavrodin (1965) and V. Sapunov (1965). 


of lion in southern Russian steppes or even the month of the Don, as 
suggested, cannot be accepted.^ 

In the Pleistocene and Holocene lions were dispersed in Europe far 
more widely than in the historic past. There is a great deal of information 
about the finds of leonine remains in southern and central Europe, France, 
England, Germany, and Poland. Such remains have been reported in the 
Soviet Union from a large number of sites in the Trans-Caucasus and Crimea 
as far as Yaroslav and Molotov regions, the Urals, and Kjasnoyar.^^ At 
the same time it is extremely difficult to study the distribution of lion in 
that period since the identification of big cats is difficult and lion is 
sometimes mistaken for tiger, especially in the case of the extinct tiger- 
lion (cave lion, P. speled). This is true, for example, with regard to specimens 
from Chernigov and others (N.K. Vereshchagin) or the Polish lion, now 
regarded as P. spelea (Koval'skii, 1959). Nevertheless, some researchers 
(Hemmer, 1967) consider the big Pleistocene cats lions, and the origin of 
the species is placed in the Early Pleistocene. 

Geographic Range outside the Soviet Union 

The range in Asia and Europe, which in part includes the Trans-Caucasian 
section of the habitation of lion, and some stages of the disappearance of 
these animals are as follows. In Europe during the historic period lions 
inhabited the southern part of the Balkan Peninsula in the north up to at 
80 least Thrace and Macedonia and probably even the Danube. Their disap- 
pearance in Greece occurred at the end of the first century A.D. (Harper, 
1945). In Asia Minor the animals were widely dispersed and evidently absent 
only in the northern part covered by the Pontic mountains (Hemmer, 1967). 
They disappeared from the western parts of the country [Turkey] evidently 
much earlier; in the east they survived up to the nineteenth century in regions 
adjoining Mesopotamia and Syria and disappeared from there only around 
the middle of that century. In the early 1870's they were still sighted in 
the upper reaches of Euphrates (Biledzhik; Danford and Alston, 1877 and 

^^Remains of lions from Ol'via (mouth of the Bug) belong to the fifth to the second 
century B.C. and are those of animals from a menagerie; some could well be only bones 
or skins of animals used as rags. These "remnants" cannot serve as proof of the occurrence 
of wild lions in Ol'via (Gromova, 1928; Topachevskii, 1956; Vereshchagin, 1959). One must 
keep in mind that if a distinction between lion and tiger is difficult on the basis of their 
skulls, it is impossible from the remnants of a skeleton, especially metapodia. It has been 
suggested that there are no differences between these two species in the postcranial skeleton. 
At the same time, tiger has never been mentioned in references to OlVia (see section on 
tiger here and also Heptner, 1959). (V.H.) 

^•^For a list of these finds, see I.G. Pidoplichko (1951). 


1880), but at the end of that century were no longer present. In Mesopotamia 
and Arabia lions still survived in that period. In the 1850's they reached 
the upper courses of the Tigris (Mosul) in Iraq, and in the 1860's were 
extremely abundant in reed marshes along the banks of the Tigris and 
Euphrates, mostly their lower reaches (Blanford, 1876). 

Lions also occurred in Kurdistan, Armenia, the eastern parts of 
present-day Turkey, and western Iran (Fig. 38). In Iran they were wide- 
spread, the Trans-Caucasian range being mainly linked with the Iranian 
range. In the far remote past they evidently covered a very significant 
part, if not much, of the country, and reached Teheran in the north, occupy- 
ing even the Persian upland. In the 1870's they were sighted only in the 
south along the western slopes of the Zagros mountains, adjacent to 
Mesopotamia (east of the Tigris valley), and in the forest regions to the 
south and southeast of Shiraz (Blanford, 1876). In southern Iran lions were 
encountered at places, though rarely, even around 900 AD. (Mekran 
coast).^^ The last of the species was killed in Iran in 1930 (Haltenorth and 
Trenze, 1956). There is every reason to believe that the "king of the jungle" 

prowled even in Afghanistan, at least in the southwestern and southern 
areas. ^2 

In India the range in the past covered the whole northern sector 
from Sind in the west to Bengal in the east and from the Narmada River 
in south (somewhat south of the tropics) to Rampur and Rohilkhand 
(about 20°30' NL) in the north (Pocock, 1939). Lions have disappeared 
throughout this territory except for a small section in the Gir forest in 
Kathiawar (see Fig. 38) where they have long been protected. These 
animals are the sole survivors of the Asian lion which, in the past, was 
not only widely distributed but, in some countries, for example Syria 
during the Crusades (eleventh to the thirteenth centuries), quite abundant 
(Usama ibn-Munkyz, 1922). 

Apart from Europe and the Near East, the range covers all of Africa 
except regions of humid tropical forests in western Africa. By the twentieth 
century the range had shrunk and lions had become extinct in the southern 
parts of Africa and in much of the northern mainland. In the 1950's (see 
Fig. 38) the animal was present only in tropical Africa from Somalia, Sudan, 
and Senegal southward to southwestern Africa, Kruger Park in Transvaal, 
and possibly Zulu- and Swaziland. (V.H.) 

■'^Personal communication from preparators NA. Zarudnyi and SA. Aleksandrov. (V.H.) 
^^The occurrence of lion in Afghanistan in the past is sometimes doubted. It is inferred 

here from the nature of the range but direct references to lion in this country are also available. 

Pocock (1939) assumed the existence of lions in the southern parts of Afghanistan where, 

according to some data (Harper, 1945), this animal was actually sighted in the middle of 

the last century. 


81 Fig. 38. Reconstructed species range of lion, Panthera leo L. Broken line 

denotes roughly the northern and southern boundaries of distribution by the 
1950's. Dot indicates present habitation of lion in India, in Gir, Kathiawar. V.G. 


Geographic Variation 

A large number of forms have been described. Of these, eleven are usually 
recognized: ten in Africa and one in Asian part of the range. Of the African 
forms, two are totally extinct and one almost so, and preserved only in 
sanctuaries (see below). 

There is [was] only one form in the Soviet Union. 

Persian lion, P. I. persica Meyer, 1826. 

This lion is wholly similar to the African forms but differs generally in 

the less swollen tympanic bullae, the much shorter postorbital constriction, 

81 and usually by the divided (doubled) infraorbital foramen (Pocock, 1939). 

The color varies from reddish-brown and highly mottled black to sandy 
or cinnamon-gray. The mane in males varies in growth and color, being 
sometimes small but usually dense on the head, neck, and chest, as also 
on the abdomen and flanks. Ulnar hair is well developed. Usually mane 
hair is rusty-brown with an admixture of blackish and white hair; it may 
also be totally golden-brown or contain a goodly admixture of black hair 
along the top and chest. Ulnar hair and the abdominal and lateral manes are 



relatively more dense than the mane proper.^^ The tail tassel is large 
(Pocock, 1939; Hemraer, 1967; Fig. 39). 

In overall size the Persian lion corresponds to central African animals. 
Body length of adult males about 195 cm (two animals) and tail length 
about 80 to 90 cm (two animals). 

Maximum skull length in aduh males (three) 330 to 340 mm and 
in females (three) 292 to 302 mm; condylobasal length in males 299 
to 313 mm and in females 266 to 277 mm; zygomatic width in males 
221 to 231 mm and in females 195 to 209 mm (data for Gir lions; Pocock, 

This species used to exist in Trans-Caucasus (up to the tenth century 

Outside the Soviet Union this species used to live in Greece, Asia Minor, 
Syria, Palestine, the Arabian peninsula, Iraq, Iran, Afghanistan, and Pakistan, 
and presently lives in the Kathiawar peninsula of India. 

Evidently lions were not entirely identical throughout the above territory. 
In Greece and Asia Minor animals with a less developed mane, without 
abdominal and lateral manes, and without ulnar hair predominated. In Syria 
and countries on the east coast of the Mediterranean Sea, and possibly the 
Arabian peninsula, lions of this type also lived, which were very similar 
and/or identical to the Egyptian lion (P. /. leo Hemmer, 1963).^"^ There 

82 Fig. 39. Main types of manes of Middle East (Persian) lion, Panthera leopersica 

Meyer according to Hemmer, 1967 (mane hair shown in black only for purposes 
of visual effect). Sketch by N.N. Kondakov. 

^■^In literature, especially popular literature, the view prevailed that the mane of 
Persian-Indian lion was very small or totally undeveloped, or almost so. Descriptions of 
maneless lions of Gujarat are available. This misunderstanding is based on the title of an 
article published in 1833 by Officer Smi, one of the first to describe Indian lions. Compared 
with lions of the Atlas mountains and menageries in which the mane is large, the mane of 
Gujarat (Gir) animals appeared small to him; he used the unfortunate expression "maneless," 
which has come into wide usage (for more details, see Pocock, 1939). 

^"'Based on a study of extensive material, i.e., ancient Greek, Hittite, Aramaic, 
Urartaean, Assyrian, and Babylonian works of art, H. Hemmer (1963, 1966, and 1967) 
reconstructed the external appearance of lions of Europe and the Middle East. 


were all types of transitional forms to the above-described Persian lion, 
which was the lone or predominant form in Kurdistan, and apparently in 
the Trans-Caucasus (Hemmer, 1966, and 1967). 

The following forms are usually recognized for Africa: 1) Berber lion, P. I. 
leo Linn., 1758 — North Africa from Atlas to Egypt (extinct); 2) Senegal 
lion, P. L senegalensis Meyer, 1826 — from Senegal to Nigeria; 3) Cameroon 
lion. P.. I. kamptzi Matschie, 1900 — northern Cameroon and the region south 
of Lake Chad; 4) Congolese lion, P. I. azandica Allen, 1924 — northeast 
Belgium Congo; 5) Masai lion, P. /, massaica Neumann, 1900 — Ethiopia, 
Kenya, and Tanganyika up to Mozambique; 6) Uganda lion, P. I. nyanzae 
Heller, 1913 — Uganda; 7) Katanga lion, P. /. bleyenberghi Lonnberg, 
1914 — Katanga, Angola, and Rhodesia; 8) Transvaal lion, P. I. krugeri 
Roberts, 1929 — Transvaal and Orange Free State; 9) Kalahari lion, P. I. 
vernayi Roberts, 1948 — Kalahari (Botswana), southwest Africa, and 
southern Angola; 10) Cape lion, P. I. melaiwchaita H. Smith, 1858 — Cape 
Colony to Natal (extinct); and 11) P. I. hoUisten Allen, 1924 — northern 
bank of Lake Victoria. 

Most of these forms have been described from extremely variable and 
insignificant characteristics, and hence their status is dubious. It has been 
suggested that all African lions are identical in subspecific features. 


There is no direct biological information about the Trans-Caucasian lion. 
This Hon lived in an environment different than the one prevailing at present 
and only under those conditions was its existence possible. The country was 
thinly populated and uncultivated or, in the Middle Ages, cultivated only 
83 here and there. At the same time the population of domestic cattle, on which 
the animal depends for survival at many places even today (Gir and some 
regions of Africa) was large in the remote past in Trans-Caucasus. Wild 
ungulates were additionally abundant. Not only large rivers like Kur' and 
Araks, but also small streams had rich forests along their banks; dense reed 
thickets grew along the extension of the Mugansk steppe and at other places 
in the lowlands; pistachio and juniper forests covered the foothills and there 
were rich pastures. All these supported a large number of deer, roe deer, 
wild boar, while in the steppes were large numbers of goitered gazelle and 
kulan. Huge reser\'es of ungulates were available to lions as food (cheetah 


also fed on them). The disappearance of lion is naturally associated with 
an increase in human population and a change in environmental conditions, 
which in turn led to an impoverishment of ungulates in the country; 
thus, kulan had disappeared from the Trans -Caucasus in the thirteenth 

Indubitably, however, direct killing also played a significant role. Big 
predators in general are few and hence easily exposed to hunting. Further- 
more, lions from the time of the ancient kingdoms of Egypt and Mesopotamia 
represented a favorite object of hunting by emperors, pharaohs, and feudal 
kings. Lions were also hunted by the shirvans of Trans-Caucasus who, of 
course, were not the only ones responsible for their being killed 
(Vereshchagin, 1949). 

At present, this subspecies is preserved only in the Gir preserve in 
Kathiawar peninsula (northwest India); the population is very small and 
rapidly declining in spite of protection. By 1960 the lion population according 
to official information was 283, but actually there were probably not more 
than 150 (Ullbrich, 1962); in 1963 they were placed at 285 but by June, 
1968 only 162 remained {lUCN Bulletin, no. 8, p. 61, 1968). (V.H.) 


Panthera tigris Linnaeus, 1758 

1758. Felis tigris. Linnaeus. Syst. Nat., 10th ed., vol. 1, p. 41. Bengal. 
1815. Felis virgata. \\\\g&r.Abh. K.Akad. Wiss. Berlin, p. 98. Mazanderan, 

northern Iran. 
1842. Felis mongolica. Lesson. Nouv. Tabl. RegneAnim., p. 50, Bukhara, 

Mongolia, Kirghizian steppes. Nomen praeoccupatum. 
1844.^-'' (1845 and 1847). Felis tigris altaicus. Temminck. Fauna Japonica. 

Mammalia, p. 43. Korea.^^ 
1858. Tigris stiiatiis. Severtsov. Rev. Mag. ZooL, vol. 10, p. 386. Renaming 

of Felis tigris Linnaeus, 1758. 
1867. Tigris regalis Gray. Proc. Zool. Soc. London, p. 263. Renaming of 

Felis tigris Linnaeus, 1758. 

-^•''The year of description of this form was originally placed as 1845 (Brongersma, 
1935; cited by Mazak, 1965) and later corrected to 1844 (Mazak, 1967). Some specimens 
from the collection of Temminck (examples in the library of V.G. Heptner) bear the date 
1847. (V.H.) 

''^More precisely, Pisihan mountains roughly at 43° NL. This parallel traverses slightly 
south of Vladivostok, through the extreme northern part of the Korean People's Democratic 
Republic and the southern part of northeastern China (Manchuria), i.e., north of the Korean 
Peninsula. (V.H.) 


1868. Felis longlpiUs. Fitzinger. Sitz. Ber. Akad. Wiss. Wien, vol. 58, 

p. 435. Amur region.^^ 
1871. Felis tigris var. amurensis. Dode. Proc. Zool. Soc. London, p. 480. 

Ussuri estuary, Khabarovsk.^^ 
84 1904. Fells (Tigris) septentrionalis. Satunin. Zh. Priroda i Okhota, no. 7, 

p. 5. Twelve versts west of Lenkoran, eastern Trans-Caucasus. 
1904. Felis tigris coreensis. Brass. Nutzbare Tiere Ostasiens, p. 4. North 


1915. Tigris mikadoi. Satunin. Zh. Nasha Okhota, no. 7, p. 17. Korea. 

1916. Felis tigris lecoqui. Schwarz. Zool. Am., vol. 47, p. 351. Lobnor, Kurla. 
1916. Felis tigris trabata. Schwarz. Ibid., vol. 47, p. 353. Hi valley south 

of [Lake] Balkhash. 
1925. Felis tigris mandshurica. Baikov. Manchurian Tiger, p. 3, Harbin, 

northeastern China (Manchuria). 
1929. Tigris mandjurica. Baikov. Zh. Okhotnik, no. 4, p. 16. Substitute 

for mandshurica Baikoff,* 1925. 
1929. T. koraiensis (micadoi) Baikoff (ex Sat.). Baikov. Zh. Okhotnik, no. 

4, p. 16. Northeastern China (Manchuria). Substitute for koreensis 

Brass, 1904 and mikadoi Satunin, 1915. 
1935. Tigris tigris virgata Matschie (1897 ex Illiger, 1804 to 1811). Ognev. 

Zveri SSSR, vol. 3, p. 273. Substitute for virgata Illiger, 1815. (V.H.) 


Color reddish-msty or rusty-yellow with narrow black traasveise stripes. Body 
length not less than 150 cm, condylobasal length of skull 250 mm, zygomatic 
width 180 mm, and length of upper camassial tooth over 26 mm. (V.H.) 


Size very large; largest species of the family. 

In general build the tiger is wholly typical of cats and has an extended 

■^^Locality fixed by S.I. Ognev (1935) and adopted by other researchers. S.U. Stroganov 
(1962), based on the fact that Fitzinger first indicated "Middle Asia", placed Lobnor Basin 
as the type locality. 

■^^According to S.U. Stioganov (1962). This work also includes a discussion of the 
names of Russian tigei's since 1868 from a nomenclatural point of view. 

'"Spelling of author's name in taxonoraic divisions is sometimes at variance with spelling 
in text and bibliography since the Israeli orthography has been followed in this translation. 

■'^In the "Description" and "Geographic Variation" sections the works of S.I. Ognev 
(1935), S.U. Stroganov (1962), and V. Mazak (1965 and 1967) have been incorporated, in 
addition to works cited in the text. 

The characteristics given under "Description" have been supplemented and elaborated 
in "Geographic Variation". 


86 Fig. 41 . Amur tiger, Panthera tigris altaica Temm. Moscow Zoological Garden. Photograph 

by A.P. Zhandarmov. 

supple body standing on rather short legs with a fairly long tail (Fig. 40). 
The front portion of the body is better developed than the rear and the animal 
stands higher at the shoulders than at the sacrum. This gives the impression 
of some heaviness but simultaneously of immense strength, which is 
emphasized by the powerful legs with large and broad front paws, heavy 
and invariably slightly drooping head, high withers, and prominent strong 
musculature, especially in the anterior part of the body, primarily in the 
form of powerful shoulders. The anirrial is quite slow on its feet and even 
its more rapid movements, leaps for example, appear leisurely. The same 
impression of quiet strength comes across even when the tiger is lying down. 
The general appearance of tiger is that of a huge physical force and quiet 
confidence, combined with a rather heavy grace (Fig. 41). 

Head with markedly projecting facial portion and bulging forehead. Ears 
short with a rounded, blunt apex and wide-set. Vibrissae set in four or five 
rows, white, sometimes brownish at the base, extremely resilient, up to 165 


mm long, and up to 1.5 mm thick. Pupil round and iris yellow. Claws very 
powerful, 511аф1у curved, and a pale waxy color. Their length along the 
outer arc reaches 80 or 100 mm and height of base up to 40 mm. 

Summer coat rather short, quite sparse, of uniform length throughout 
the body, with long hair present only on cheeks. Some long hair is also seen 
along the upper part of the neck. Coat quite coarse. In southern forms living 
outside the Soviet Union there is no difference between the winter and 
87 summer coats, or very little. Among tigers of Turkestan and the Caucasus, 
but especially among those of the Soviet Far East, coat differences are 
very shaф: winter coat is denser and longer, softer and silky, and might 
even appear shaggy on the trunk of animals with an especially rich coat. 
Hair is markedly longer on the head (the ears project insignificantly); 
whiskers and hair on the occiput and along the top of the neck also 
greatly elongated. 

Compared with other Soviet cats, the pelage of the tiger is of moderate 
thickness, coarse, and sparse. Back with 2,500 hairs per 1.0 cm and abdomen 
with 600; 1.4 underfur hairs occur for every 1.0 guard hairs (an extreme 
case among Soviet cats). Underfur very slightly curled and coarse. 
Exclusively black hairs (of black stripes) differ greatly from yellow hairs, 
i.e., those devoid of black pigment. Average length of yellow hair 27 mm 
and thickness 105 microns and for black hair 48 mm and 88 microns 
respectively. Length of yellow guard hair (categories I to IV) 31, 29, 25, 
and 25 mm and thickness 85, 62, 49, and 37 microns; length of black guard 
hair 40, 38, 33, and 31 mm and thickness 87, 63, 47, and 34 microns. Yellow 
underfur 24 mm long and 30 microns thick and black 23 mm and 30 microns 
respectively (winter skin of Amur tiger; B.F. Tserevitinov; also see 
"Geographic Variation" below). 

Main background color bright reddish-rust or ocherous-rust. This shade 
is brightest on the back and less vivid on the flanks and outer side of the 
legs. Lower part of head, neck, midportion of chest, abdomen, and inner 
side of legs white or with light yellowish tinge. Transition of color from 
upper to lower surface gradual but distinct. Main color of tail on dorsal 
surface same as on trunk, but toward the end slightly lighter, and may even 
become white; underside of tail white or very light ocher. 

Against this main background color an extremely complex pattern of 
dark (pure black or brownish) stripes is distinct. These .stripes vary in size, 
form, and disposition in different parts on the body; vividness and sharpness 
of outline also vary. There are no annular spots on the coat. 

The following disposition of dark stripes is typical: transverse (vertical) 
position throughout the body and along the neck; on the trunk some descend 
low while others terminate above, or even extend onto the abdomen. All 
stripes with shaф lower ends, sometimes bifurcate, or divide and fase again 


toward the end. Other transverse stripes commence between these tapered 
ends and descend onto the abdomen or encircle it. Those commencing on 
top of the neck terminate on the neck or run onto the chest; those from 
the withers descend along or onto the rear of the shoulder blades. 

Stripes are usually more densely distributed on the posterior half of 
the trunk than on the anterior. Anteriorly, however, the color is usually 
more intense — more or less pure black — and not uncommonly a brownish 
shade posteriorly; they may even be brownish on the thigh. Stripes 
commencing on the sacrum run onto the groins or to the rear of the thigh, 
or terminate midway on the thigh. Some, having reached the upper part 
of the coat, take an oblique direction, incline forward, and run onto the 
inner surface of the legs. 

The outer and inner surfaces of the hind legs up to the calcaneal joint 
are covered with dark transverse (horizontal) stripes, which are short or 
terminate in a semicircle or even a ring. Foot dirty gray below. Front (outer) 
surface of forelegs devoid of black stripes, while inner surface carries short, 
intermittent black transverse stripes, their ends sometimes extending onto 
rear (outer) surface of legs. Lower ends of stripes on shoulders and in 
scapular region may descend to the ulna. 

Tail invariably with a black tip and for much of its length covered 
\\dth transverse bands forming compact rings. Such rings are absent, however, 
at the base of the tail; dark markings occur along the top of the tail in 
the form of angles with apices turned forward. There are usually 10 dark 
transverse stripes on the tail but these may number 8, 9, or 11. 

The pattern on the head is complex (Fig. 42). Dorsal side of muzzle, 
commencing from nostrils and up to region between the eyes and slightly 
posterior, rusty-colored and without dark pattem. Region from the comer 
of the eye to upper row of vibrissae similarly colored. This coloration usually 
extends in the form of a small wedge under the eye to its outer corner. 
A narrow white field runs directly along the lower eyelid. Region of lower 
nostrils and vibrissae, chin, and lower jaw white except for black spots 
on lower lip in the corner of the mouth. Sinciput and occiput with complex 
and variable pattem of short transverse black stripes, often irregular in form; 
there are usually four, five, or six such stripes. Sometimes some or all are 
interrupted in the middle; sometimes a longitudinal stripe occurs between 
them and transverse stripes are set off from it. The last pair of transverse 
stripes sometimes forms an angle.'*^ 

''^he head pattern consisting of a longitudinal and three transverse stripes (see Fig. 
91) forms a figure which is extremely similar to the Chinese hieroglyph "van" or "van- 
da"' which means "great ruler". Among the Chinese and Koreans and some smaller tribes 
of the Far East, tigers were associated with numerous superstitions and animals with the 
sign "van" considered sacred and treated respectfully (Baikov, 1925). 


Fig. 42. Amur tiger, Panthera tigris altaica Temm. Moscow Zoological Garden, 
1%3. Photograph by V.E. Prisyazhnyuk. 

Often a long stripe begins behind the ear and descends in the form 
of an arc onto the neck, often reaching the throat. The sides of the head 
and the whiskers around the white field are covered with some black stripes 
of angular or bracket shapes. At their top they commence under the ear 
or before it. Sometimes one of the transverse bands of the sincipital pattern 
also extends down along the cheeks and flanks, reaching the corner of the 
lower jaw or even descending onto the throat. The front surface of the ear 
is covered with white hair while the rear is invariably pure black with a 
large bright white spot in the upper half.'*^ 

The above description of tiger coloration is highly schematic (see also 
"Geographic Variation") and undergoes a good deal of variation, primarily 
individual; color shade as well as pattem undergo changes. The general color 
of the coat is sometimes quite vivid and bright rusty and sometimes a dull 
yellow. The downward extension of the rusty field also varies. So too do 
the white fields on the underbody, rising to various degrees on the flanks 

''^This spot, developed in some other large (leopard) as well as small (Amur 
[= leopard] cat) cats, is of significance as a signal. 


and neck. The coloration of the pattern itself likewise varies. The black 
stripes on the trunk may be brown-toned while those on the thighs and 
rear of the paws, usually somewhat brownish, may be more vivid. 

The black pattern is subject to considerable variability. Stripes may 
be wide, narrow, continuous or bifurcate at the end, or with a light-colored 
interruption, shorter or longer, several or many, the skin densely or sparsely 
covered witlithem, and so on. Stripes on the flanks usually broaden in the 
middle of the back and, converging with the stripes from the opposite side, 
merge or alternate with them; sometimes a black longitudinal band forms 
on the back, often extending the entire length of the trunk. Sincipital pattern 
and that in general on the head, side of the muzzle, and around the eyes 
varies notably. 

Pattern of tail likewise is highly variable. In addition to 'distinct and 
well-demarcated closed rings throughout the tail one also sees tails with 
only a black tip and some distinct distal rings, or tails with paired dark 
rings and yellow bands. More often the front rings are not distinct — neither 
completely covering the tail, irregular in form, and without a distinguishable 
pattern. By and large these "rings" are angular bands distorted to various 
degrees, but with the apex turned forward. Nevertheless, whatever the 
variation, ten dark fields are invariably distinguishable. The background 
color of the tail may be white throughout its length or for most of it. 

Tigers showing sharp deviations are known. Some have a reduced 
number of black stripes in the front half of the body. In extreme cases the 
scapular region and chest are almost devoid of a black pattern, while in the 
90 rear portion of the body stripes may be normally developed. Melanism is 
known, as is albinism (some faded traces of stripes are usually discernible), 
and so are referred to as white tigers. In the latter the rust-colored pigment 
is reduced but the black pattern fully developed. These mutations have not 
been seen among Soviet animals; a reduction in number of stripes has been 
observed very rarely. 

Seasonal color variations in northern, i.e., Soviet, tigers are significant. 
The main background coat color in the winter, compared with the summer 
coat, is less bright and rusty, in other words more ocherous. As a resuh 
of the greater length of the pelage, the stripes appear broader and have less 
sharp outlines. 

Sexual dimorphism in color is not apparent. Whiskers in the winter 
coat are not so well developed in females as in old males. Age-related color 
changes are absent; young cubs have the same striped coat as adults but 
the main shade of their coat is less vivid and more ocherous. Moreover, 
the stripes in cubs are distributed closer together, are wider, and their outlines 
less sharp, due partly to the relatively long hair and furriness of the coat. 
A black line on the back is usually present. 


Geographic variation in color and pelage type is clearly expressed and 
reflected in the intensity of the rust coloration; the extent of development 
of the stripes (their number, density, width, and length), intensity of black 
coloration, size of the white field on the underside of the body, the black 
pattern on the tail, and other details of color also undergo change to some 
extent or another. The degree of seasonal dimoфhism in length and richness 
of the coat also undergoes geographic changes. The geographic color changes 
follow Gloger's rule (more details later). 

Albinism and melanism are known among tigers living in the wild. 
Under certain light conditions traces of dark stripes are usually seen among 
albinos. White tigers, present mainly in America, are those raised in 
zoological gardens from a lineage of animals in which the yellow pigment 
has failed to develop but the black pattern has remained. These animals 
owe their origin to the tiger indigenous to India. 

Teats; two pairs, inguinal. 

Diploid number of chromosomes, 38. 

The skull (Fig. 43) is wholly typical of the genus and characterized 
by its large size. In this respect, as also in almost all others, it is similar 
to the skull of lion, from which it differs only in structural features of the 
lower jaw and relative length of nasals described above (see description 
of lion).'*2 The skull of the tiger is heavy and massive with a large and very 
powerful facial region, and very broad in the region of the canines. The 
cranium is elongated and powerfully compressed with well-developed crests, 
especially the sagittal. General skull profile bulges slightly, the highest point 
being the interorbital region. Orbits are relatively small, oval in form, extend 
to the rear, and open broadly behind. Tympanic bullae are relatively small, 
thick-walled, and flattened. Mastoid process broad, massive, and conical 
in form. Lower jaw heavy, with a broad coronoid process. Lower profile 
of jaw somewhat concave (see Fig. 36). 

Upper canine unusually well developed, broad at the base, and long 
(to 65-66 and even 70 mm along front curvature). Upper molar very 
small and sometimes absent (lost). Upper camassial tooth very large, 
with well-developed antero-intemal cusp displaying a small cone; quite 
often an additional tiny cone develops on the anterior outer edge of the 

Sexual dimorphism of the skull is evident only in the slightly reduced 
overall size in females, and in old males, intense development of the crest, 
greater compression of the postorbital region, and also of the brain case itself. 

'''^Pocock (1939) pointed out that differentiation of the skulls of lion and tiger is not 
always easy when a large number of both are to be identified. It is also known that there 
are no differences in the post-cranial skeleton of the two species. 

89 Fig. 43. Skull of Amur tiger, Panthera tigris altaica Temm. No. S 2088, collection 

of Zoological Museum, Moscow University. Ussuri region. Sketch by N.N. 




Fig. 44. Age-related changes in general features and relative size of skull of tiger. 
Left — skull of an old male, No. S 2088, Ussuri region, Bogomolov, with greatest 
skull length of 360 mm. Right — skull of young animal, No. S 58977, Ussuri region, 
between Khor and Iman Rivers, winter of 1955-1956, with greatest skull length 
of 220 mm. Collection of Zoological Museum, Moscow University. Sketch by 

N.N. Kondakov. 

Age-related changes in the skull are very significant (Fig. 44). The 
skull of young animals is rounded with relatively poorly developed and less 
massive facial region, a more bulging and relatively large brain case, and 
91 zygomatic arches which are weaker, not so angular, and bulge less. With 
age the skull elongates; the facial region develops at the expense of the 
cranial part; and the skull becomes heavier, thicker- walled, and uneven. 
On the whole the range of age-related skull changes is great, the greatest 
in the family (as in lion). Geographic variability in skull structure is small 
and pertains only to some differences in overall size. 


The tiger is the largest species of the family, its body size and weight 
being more than that of the lion. At the same time the largest skulls of 
lion are only slightly bigger than the largest skulls of tiger."*^ The lion 
evidently has a somewhat larger head and a shorter body, which is generally 
less massive.'*'^ In size and weight the tigress is much smaller than the 
male. Differences in old animals may reach 15 to 20% in body length and 
up to 50% in weight. By and large, however, size in these animals is 
extremely variable; this variability resuhing not only from the common 
individual variations, but also from the fact that tigers live long, continue 
to grow for a long period (males almost throughout their life), and under 
certain conditions may become extremely plump. The absolute body and 
skull size and weight of Soviet tigers, which provide an idea of the species, 
are given under "Geographic Variation". 
92 The geographic variability in body and skull size is distinctly manifested 

and follows Bergmann's rule, but not rigidly. The dimensions of some 
intemal organs based on the measurement of three wild adult tigresses follow 
(No. S 34885, Ussuri region. Zoological Museum of Moscow University, 
body length 172 cm, weight 145 kg; No. 197, Pyandzha, Tadzhikistan, body 
length 165 cm, weight 135 kg; and No. 203, Kyzyl Su, Tadzhikistan, body 
length 165 cm, weight 97 kg; data of Chemyshev, 1958, with modificatioas, 
and V.G. Heptner): weight of heart and heart index 770 g/5.28; 570 g/4.22; 
and 970 g/10.00 (V.H.); weight of lungs 850, 1,480, and 2,070 g; weight 
of liver 1,720, 1,955 and 2,179 g; weight of spleen 265, 290, and 250 g; 
weight of alimentar}' canal 2,500 and 2,700 g; weight of stomach 795 and 
910 g; weight of kidneys 990 and 1,160 g; total length of intestines and 
ratio to body length 742 (1 : 4.31), 640 (1 : 3.88), and 710 cm (1 : 4.30); 
length of small intestine 542 cm (87.5% of total length of intestines), 550 
(85T9%), and 660 cm (92.9%); length of large intestine 100 (13.5%), 90 
(13.4%), and 50 cm (5.6%); and length of appendix 6.0 cm (0.8%), 5.0 cm 
(0.7%), and 10.5 cm (1.5%). Cross section of heart 118 mm and length 
132 mm (specimens of the Zoological Museum of Moscow University). 

Systematic Position 

In spite of considerable differences in the general appearance between lion 
and tiger, these two species should be regarded as very close to each other. 
The differences in their skulls, as shown above, are very insignificant and 

''^he maximum skull length of tiger is evidently equal to 385 mm (V.G. Heptner; see 
later) or possibly 400 mm (Baikov, 1925). Among lions the maximum skull length may reach 
390 and even 400 mm (Roberts, 1962), the world record being 419 mm (Best et al., 1962). 

'*''lt has been shown (Best et al., 1962) that in India, the lion is "inferior" to the tiger. 


; Fig. 45. Amur tigress aged 3.5 to 4.0 years with relatively shaф and broad stripes 

in anterior body and narrower and closer set stripes in rear. Body length 159 cm, 
tail length 88 cm, and height at withers 79 cm. December 11, 1961, Suchan River, 
southern Ussuri region. Photograph by V.K. Abramov. 

often inconstant; the same is true of differences in the postcranial skeleton. 
While some subspecies of lion and tiger differ greatly, at the same time 
resemblances, for example between tigers and lions of Asia, are quite 
significant. This fact has provided a basis for some zoologists to place the 
origin of these cats in southwest Asia. On the whole the differences between 
these two species, taken in toto, are less in craniological features than between 
other proximate species of cats. Most systematists place these two species 
in immediate proximity in taxonomic tables. 

Yet despite the foregoing, Hemmer (1966) has expressed the view that 
tiger is a distinctive monotypic subgenus and stands in contrast to all other 
big cats, which constitute a different subgenus. Furthermore the lion consti- 
tutes a species distant from tiger but most closely linked with leopard, and 
93 through it with jaguar. In such a sequence the ethological peculiarities 
of the species, as also the much easier hybridization of lion with leopard 
and jaguar than with tiger play an important role. There is no justification 
at present to adopt this scheme; it runs contrary to morphology. (V.H.) 

Geographic Distribution 

Tigers live in forests and reed jungles in southern, southeastern, and Middle 
Asia, and parts of the Near East and Central Asia. 


Geographic Range in the Soviet Union 

The range in the Soviet Union (reconstructed) constitutes the northern rim 
of distribution of the species. Within the Soviet Union the range is divided 
into three independent sections — Caucasian (including the European part 
of the USSR), Middle Asian (including Kazakhstan and western Siberia), 
and Far Eastern (including Trans-Baikal). These areas join in the south out- 
side the USSR. The Middle Asian, and part of the Far Eastern range, actually 
consists of isolated sections or pockets (Fig. 46). 

In view of the already mentioned peripheral nature of the range in the 
Soviet Union, two parts have been identified, or are being identified, in 
each of the three habitations of tiger in this country; these are territories 
of permanent habitation and breeding, and areas where the animals are simply 
intruders. Usually the region of intrusion adjoins that of a somewhat wider 
zone of permanent habitation. In relation to the entire range of the species, 
the area of transgression in different parts of the periphery cannot be very 
large. However, given the relatively small sections of the range falling within 
the Soviet Union, this area is quite significant. At places the area of 
transgression could conceivably exceed the area of permanent habitation. 
One must also keep in mind that the separation of these two zones is not 
always possible; thus the true boundary of permanent habitation of the 
species can hardly be delimited when discussing the past or the period of 
disappearance of the animal from any part of the range when its normal 
population and life pattern were disturbed. Tigers are highly mobile and 
intrusions into other places are frequent. They may be seen for several years 
almost regularly in a certain area, or for some period become residents and 
even breed, then for some years disappear altogether, or occur as rare 
intmders. Sometimes the boundaries shift in a discemible direction. Finally, 
in some wide regions in the past the animals settled, but later under the 
impact of extinction appeared as intruders. 

Caucasus. In the nineteenth century (Fig. 47), the permanent habitation 
and evidently also breeding grounds of the tiger covered only the extreme 
southeastern part of the country — Talysh and Lenkoran lowlands. Here 
tigers were sighted in hilly forests up to the northern extremity of the 
mountains and in lowland forests. They were encountered most often in 
the lowland forests of Prishib. Here their population was quite large until 
the end of the last century and even early in the present century, although 
significant fluctuations were observed at different periods. 

A rapid decrease in the number of animals was recorded for the early 
years of the present century and even the end of the last. The animals stopped 
breeding and were found only as regular intruders from Iran that remained 
in the Caucasus for quite some time. This was primarily due to direct 

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persecution by the expanding Russian populace, changes in topography 
(deforestation and cultivation of the land), and extinction of wild boar — 
the main prey of tiger. 

In the last century, and somewhat even before, when Talysh was a 
permanent habitation for tiger, the animals moved out from there into 
different parts of the Caucasus, mostly the plains of the eastern Trans- 
Caucasus. Talysh later became the site of intrusion of tiger into Russia from 
Iran. This is because Talysh lies in the westernmost extremity and is covered 
with the forested landscape of the Cis-Caspian province of Iran that is 
connected to Elburz (Hyrcania). Evidently only a few tiger intrusions into 
the Caucasus were not linked with Talysh, occurring instead through the 
more western parts of northwestern Iran (Zangezur range at Araks in the 
region of Megra, Armenia; Ararat depression; probably, Akhaltsikha). From 

Fig. 47. Northern boundary of region of permanent rangeof tiger in the Caucasus 

in the nineteenth century (1); individual sites (2); and region of intrusion in the 

eighteenth, nineteenth, and twentieth centuries (3). V.G. Heptner. 


Talysh the animals most often intmded into the nearest areas of Trans- 
Caucasus steppe (Belyasuvar in Mugansk steppe, Sal'yana, and others) and 
also penetrated significantly farther, to Tbilisi, reaching there evidently along 
the Kur' valley (1735, 1820, and 1922). Tiger intrusions from Iran into Talysh 
have become quite rare in the last decade. Reports of such came in the 1950's 
and 1960's but the last (to 1969) was recorded in 1966 (Lenkoran low- 
land).'^-'' The appearance of stray animals even now is not excluded, but 
its possibility is very remote since the population is rapidly declining in 
Hyrcania. By the middle of the twentieth century tiger could be considered 
extinct in the Caucasus. "^^ 

In the eighteenth century and the first half of the nineteenth tiger 
transgressions beyond Talysh were far more numerous and covered the north, 
east and west more extensively. In the north the animals reached Shemakha 
(foothills) and Baku and even Derbent; in the east they were sighted in 
Armenia and Georgia proper (in the east — region of Tbilisi, 1820 and 1835); 
and in the western Trans-Caucasus in Kvirila and Rioni basins (Imeretiya 
and Mingreliya), including the southern slopes of the Great Caucasus [range]. 
The northwestemmost section of the range boundary of the species evidently 
traversed the latter (Brandt, 1856).'*'' The boundaries of permanent hab- 
96 itation and region of intrusions are not known. It is quite clear, however, 
that Armenia and westem Trans-Caucasus, like Tbilisi, constituted regions 
of intrusions. At the same time, probably in the eighteenth century, the 
animals lived permanently not only in Talysh and Lenkoran lowlands 
but also along tugais, rushes, and lower reaches of the Kur' and Araks 

There is justification for believing that in the Middle Ages, especially 
in the tenth to the twelfth centuries, perhaps even later, tigers lived perma- 
nently at suitable sites throughout the plains of eastern Trans-Caucasus and, 
possibly, at some places in the foothills of the Great and Little Caucasus 
(Fig. 48). 

The natural environment in these regions was far more favorable for 
tiger in the Middle Ages than in the nineteenth century. Humans were few, 
water sources full, tugais and all types of trees abundant along rivers and 

''■''Some additional infomiation about tiger intiusions into the Caucasus and other parts 
of the range may also be found in the section "Biology" ("Population" and "Seasonal 
Migrations and Transgressions"). 

''^Brandt, 1956; Satunin, 1903 and 1915; Dinnik, 1914; Smirnov, 1922; Ognev. 1935; 
Vereshchagin, 1947 and 1959; Sludskii, 1953a and 1966. 

''^Sometimes the data of Giildenstaedt (1787 and 1791), on which Brandt relied, pertain 
to leopard; these are hardly reliable data. References to tiger in western Trans-Caucasus 
are also found in other older works (Sharden, Gamba, Nordmann, and others; Brandt, 1856), 
even for the mountains south of Akhaltsikha in the nineteeth century. 


lakes and, at places, even along the sea coast; wild boar, deer, roe deer, 
kulan and goitered gazelle were numerous (see section on lion). Along the 
Caspian coast the animals evidently penetrated far north. 

There is every justification for believing that the ''fierce animal" of 
Russian literature in the Middle Ages was not the wolf, leopard, or a lion, 
as presumed by many researchers, but the tiger (Heptner, 1969). It may 
therefore be assumed that tiger occupied even the northern Cis-Caucasian 
plains; not only the Cis-Caspian rushes far in the north but also Terek, 
Kuban, and Azov coasts. It colonized even the estuary of the Don and the 
southern Russian steppe, and probably even the forest steppe (Chernigov 
principality). Today one must only speculate on the actual outlines of the 
range and the nature of occurrence of the animal in this extensive expanse 
north of the Great Caucasus. It is highly probable, however, that the animal 

96 Fig. 48. Probable range of tiger in the northwestern section of the range in the 

Middle Ages (tenth to twelfth centuries). 


97 was a permanent inhabitant, at least in the northern Cis-Caucasus (for more 
details, see Heptner, 1969). 

Middle Asian area of оссштепсе. Within the Soviet Union this region 
is divided into several independent sections (Fig. 49). In southwestern 
Turkmenia tigers in the past evidently lived permanently along the Atrek 
[River] as far as the sea (Chikishlyar) and along its tributaries, Sumbar 
and Chandyr, in any case along their lower reaches. Insofar as the rest of 
Kopet-Dag within the Soviet Union is concerned, tigers were encountered 
there rather often but only as intruders. Tigers lived in southwestern 
Turkmenia during the last century, but evidently only up to the 1880's or 
probably 1890's. Their disappearance was directly linked with their 
persecution, mainly as a result of Russian colonization, and also changes 
in their natural habitat, i.e., the disappearance of rushes along Sumbar and 
Chandyr and later even along Atrek, and a reduction in population of wild 

In regions very close to the Atrek basin and along Gorgan in Iran, tigers 
survived well and at places (Talau) lived even until the middle of the present 
century. From there they penetrated, and continue to do so, into the Soviet 
Union. Transgressions occur once every several years, mainly into the 
western and southwestern parts of Kopet-Dag. The animals have been caught 
on the Atrek, especially at Kyzyl-Atrek and Chat, where four tigers were 
killed separately in April, 1930 (V.G. Heptner) — at Kara-Kara and at 
Kaine-Kasyr on the Sumbar, at Nukhur, and even at Kyzyl-Arvat, in the 
environs of Ashkhabad in the northern foothills. An intrusion is on record 
into the Great Balkhan, i.e., through the vast expanses of the desert.'*^'' In 
the 1930's intrusions were rarer than in the 1920's and earlier, and in the 

98 1940 's even rarer than in the 1930's and so on. In the eighteenth century 
tigers were reported on Cheleken Island (V.G. Heptner) and near Balkhan 
Gulf (Brandt, 1856, according to Gmelin). 

The habitation of tiger in southwestern Turkmenia is directly associated 
with the habitation of the animal along Elburz [range]. The former represents 
the northeastern extremity of the range, in the same way that Talysh 
represents the northwestern extremity. 

Further east, in the expanse between Ashkhabad and Tedzhen, tigers were 
evidently absent. This region was altogether unsuitable for these animals 
(Takyr zone at the foot of the Kopet-Dag range) and there are no data even 
about transgressions. In Tedzhen tigers were reported from Puli Khatun (35° 

''^he last tiger (to 1969) in Kopet-Dag was killed on January 10, 1954 on the Sumbar 
around Kaine-Kasyr right nn the Iranian border (Shukurov, 1958; in this work data are also 
available on other finds in Kopet-Dag). It is suggested that some confusion with leopard 
may have occuired in the reference to the Great Balkhan. However, tigers move long distances 
across the desert (see below). 


120 О 120 240 360 km 


Fig. 49. Middle Asian area of occurrence of tiger in the Soviet Union. 

1 — boundary of pennanent habitation and regular short intrusions in the nineteenth 
century; 2 — sites of stray finds at the end of the eighteenth, nineteenth, and partly 
twentieth centuries (most intrusions in the twentieth centur)' were in the region 
of former permanent habitation); 3 — migrations from Amu-Darya delta into Syr- 
Darya and vice versa, and from Chu River into Semirech'e; 4 — not wholly reliable 
information about Zeravshan and unconfirmed limit of occurrence along Sary Su. 

V.G. Heptner. 

55') up to the liver mouth (Zarudnyi, 1890 and 1891). They were undoubtedly 
encountered even beyond since they bred in Afghanistan along the upper 
reaches of Hari-Rud (i.e., Tedzhen) at Herat. Along the tugais and reed 
jungles in the lower reaches of the river at Sarakhs and somewhat higher, 
the animals undoubtedly lived permanently; they were probably only regular 
visitors in the much higher levels where large tugais and rushes could not 
have been present because of the very nature of the river valley (it cuts 
through the Gyaz'-Gyadyk mountains). 


Around Tedzhen tigers were common in the 1880 's (especially around 
Karry-Bend railway station) and were encountered even in the 1890's. 
Destruction of forests and mthless killing led to their total disappearance 
early in the present century. There is no information for the present century 
about even intrusions at Tedzhen. The Tedzhen section of tiger habitat was 
evidently connected with the animal 's occurrence in northeastern Iran, as 
well as the Afghan section of the range. 

In the past tigers were extensively distributed and at places very common 
around Murgab and the state boundary to Mara (Merv) in the north. They 
were also sighted along the Kushka from Chil'dukhtar (south of Kushka 
town) up to its confluence with the Murgab. Evidently they also occurred 
on the Kashan, which enters the Murgab above Kushka. Along the Kushka 
tigers were seen mostly as intruders though quite regularly, and could have 
existed as permanent inhabitants in some periods. The disappearance of 
tigers on the Murgab and Kushka is associated with the destruction of the 
tugais and cultivation of the valley (cotton), but more so with ruthless 
persecution, even in the pre-Revolutionary period, especially by hunting 
parties of the army and some military sportsmen-officers. 

The animal had already disappeared long before in the north, but 
persisted in the south until the 1880's and even 1890's. On the upper Kushka 
it was fairly common at that time (1880 's). By the early present century, 
tigers were practically extinct throughout the Murgab basin and there is 
no reliable information even about their intrusion into the upper Murgab 
and Kushka. The possibility of intmsions is not excluded even in the 1920's 
since the animal was still sighted though rarely, in adjacent parts of 
Afghanistan. Even before the middle of the present century, however, they 
had disappeared there."*'^ 

The Amu-Darya basin represents a more or less isolated area of 
occurrence of tiger. The animals occurred from the lower reaches of Pyandzh 
as far as the Aral Sea. Along this entire stretch two main isolated sections 
of permanent habitation of tiger occur — northern and southern. In the 
intervening region stray animals occurred and, if they stayed there at all, 
residence was brief. Tigers did not flourish, or only rarely flourished, or 
lived there permanently so long ago that no precise information is available 
about their prior occurrence. 

The northern section covers the whole deha of the Amu-Darya where 
tigers lived in the area northward up to Nukus and Khodzheila. Formerly, 
tigers were encountered in the tugais along the Amu-Darya, and significantly 

■^^According to data of NikoPskii, 1886; Radde and Walter, 1889; Eichison, 1889; 
Zai-udnyi, 1890 and 1891; Silanfev, 1898; Satunin, 1905; Masal'skii, 1913; Bil'kevich, 1918 
and 1924; Flerov, 1932; Leptev, 1934; Heptner, 1949 and 1956; Sludskii, 1953 and 1966; 
Shukurov, 1958; and others; and data of V.G. Heptner. 


farther, not only at Khivinsk oasis but also in the Dargansk Ata (40°30') 
and Dayakh-tyn (40°05') tugais. Throughout this region, particularly in the 
above two southern tugais, tigers were encountered even at the end of the 
last century and in the early present century, and were found in significant 
numbers in the delta. Destruction of the tugais, extinction of wild boar, and 
99 tiger hunting led to a rapid drop in numbers and a reduction in range. By 
the 1920's tigers were sighted only in the delta and even in the 1930's only 
around Khodzheila and Nukus. The last of the animals at Nukus was killed 
in 1938. In the 1940's they were seen only in the lower regions of the delta 
(in the region on Kungrad, Karauzyak, Muinak, and Kegeili) where some 
12 to 15 animals were counted in 1942. In spite of a ban on their hunting, 
their numbers had declined to about five or six in 1947, and these had 
disappeared by 1950.-''^ 

The southern section of occurrence of the tiger along Amu-Darya 
includes the Pyandzh valley and the uppermost reaches of the Amu-Darya 
and its right tributaries. In the east the tiger reached Chubek (south of 
Kulyab) and the western foothills of the Darvaza range. It lived permanently 
among the tugais and reed jungles of the Pyandzh and Amu-Darya as far 
as Termez, and probably lower. It lived permanently also among the tugais 
of the Yakhsu, Kyzylsu, Vaklish, Kafimigan, Surkhan-Darya, and Shirabad. 
The animal ascended very high along them — as far as the Gissar valley 
(upper reaches of Kafirnigan) to the upper reaches of the Surkhan-Darya. 
The region of permanent habitation and breeding of tigers was connected 
predominantly with the lower reaches of the above rivers; in the higher 
parts of their valleys the animals were mostly intruders. 

This region was the richest in tigers in Middle Asia. This was not only 
because tigers multiplied undisturbed under the extremely favorable condi- 
tions in the forests and the unusual abundance of wild boar and deer, but 
also because their numbers were invariably supplemented by migrants 
entering the region from Afghanistan.-''^ This was the situation prevailing 

''"^According to some data the last of the animals was killed in 1947 and information 
about killing of tigei-s in the delta in 1950 and 1951 is regarded as dubious (Sludskii, 1953). 
However, information is available concerning sightings of lone animals in the delta in 1955 
(February), in 1963 (December: Kegeili region), and in 1965 (September; Mambetzhumaev 
and Palvanniyazov, 1968). Finally, in mid-June, 1968 one tiger, evidently not old, was 
encountered on the bank of the Amu-Darya 25 km above Nukus. Probably, the very same 
animal was sighted by a large group of people in the foothills of Sultanuizdag 3.0 km away 
from the right bank of Amu-Darya, 15 to 20 km beyond the point of first sighting in June, 
1968. This suggests that tigers may still be living in several dense areas of the delta (A.M. 
Mambetzhumaev and A.S. Sabilaev). 

^Юп Urta-tugai Island (about 400 km- in area, now part of Afghanistan) on the 
Pyandzh at Parhar, some four to five litters were discovered annually according to soflie 
accounts. This is obviously an exaggeration. 


at the end of the last century and partly in the first decade of the present. 
Yet even toward the end of the last century their decline had begun. Changes 
in habitat played a decisive role only in the recent period (from the 1920's 
and especially 1930's). The main reason, however, was the extermination 
of the animal by military game hunters.-'*^ 

Even in the 1930's tigers bred along Pyandzh and Amu-Darya and in 
the lower reaches of Vakhsh and probably in other isolated places, but 
by 1950 (probably earlier) their population here consisted of only a few 
stray animals coming from the south. They lived more permanently in the 
lower reaches of Vakhsh, especially in Tigrovaya Balka preserve. In the 
1930's, 1940's, and early 1950's instances of tiger appearances were recorded 
in Surkhan-Darya valley at Denau and Saryassiya (1930; V.G. Heptner), 
the floodplains of the Kafimigan in the north as far as Mikoyanabad 
(Kabadian) in the Gissar valley (killed at Rokhatinisk MTS in 1938), the 
Vakhsh valley in the north of Aral (Kuibyshev region), the Pyandzh, 
especially its upper reaches, the lower reaches of Kyzylsu in the region 
of Parhar, and along the river below Kolkhozabad. In the Kyzylsu tugais 
by 1950 there were not more than five tigers and their total number in 
Tadzhikistan as a whole did not exceed ten to fifteen. In the sanctuary, 
"Tigrovaya Balka", tracks were seen for the last time in 1953. Thus in 
the region under description tigers had disappeared by the 1950's but some 
stray transgressions from Afghanistan were still possible. Furthermore, even 
in that country the animal was very rare along the Pyandzh and Amu- 
Darya; and at the end of the 1960's tigers reportedly entered Afghanistan 
from the Soviet Union (J. Niethammer). 

Between the above two sections of permanent habitation of tiger, along 
Amu-Darya, in a large section of the river valley the animals did not live 
100 permanently, at least not in the last 50 to 70 years, but were often sighted 
as intruders. Reports include tiger appearances at Kyzyl- Ayak (25 km south 
of Kerka), at Kerka, at Pal'vart (80 km below Kerka), and at Chardzhou. 
In some cases the animals may have stayed for some time and even 
reproduced. In the winter of 1931 a group of six tigers migrated beyond 
Chardzhou, down toward the Amu-Darya, and settled (? V. H.) at Dzhar- 
rabat-tugai (about 30 to 40 km below Darganat) and reproduced there. In 
the past, when the tugai and forests were plentiful and tigers abundant in 
the delta as well as in the upper reaches of Amu-Darya, such migrations 
along the river were more frequent.-''^ 

^^Some had in their possession several tens (!) of tiger skins (Masal'skii, 1913). 

"Bogdanov, 1882; Lagofet, 1913; Masal'skii, 1913; BiFkevich, 1924; Gladkov and 
Nikol'skii, 1935; Flerov, 1935; Sludskii, 1950, 1953, 1953a, and 1966; Chemyshev, 1950 
and 1958; Berg and Ivanova-Berg, 1951; Pokrovskii, 1951; and others; and data of 
V.G. Heptner. 


In the remote past tigers undoubtedly lived or were present on the 
Zeravshan, probably even in the middle of the last century, but references 
to this are mainly indirect (Severtsov, 1873). 

Along the Syr-Darya tigeis were found from the estuary to the Ferghana 
valley. Along the right tributary of the Syr-Darya, i.e., Chirchik, they were 
encountered as far as Tashkent (tigers lived permanently in the immediate 
proximity of the town in the 1870 's) and above (reported for the Kuraminsk 
range 40 km south of Tashkent), and noted along the Arysu and its tributaries 
the Boroldai (50 km north of Chimkent) and the Ul'kun-Aksu (in its upper 
reaches in the western spur of the Talassk Alatau). They also occurred along 
the entire coast of the Aral Sea where large reed jungles were abundant 
in the past. Tigers likewise lived in the lower reaches of the Kuvan-Darya 
and Dzhany-Darya. 

Tigers were widely distributed in Syr-Darya basin in the early and 
middle parts of the last century, although even then at some places, for 
example in the Ferghana valley, they were reported only as intruders and 
evidently rare. The extinction of tigers became more rapid with the 
appearance of Russian hunters, their population began to dwindle rapidly, 
and their area of distribution shrank. Tigers survived the longest in the lower 
reaches of Syr-Darya below Kazalinsk. Nevertheless, at the end of the 
nineteenth century they were evidently still met with along the river and 
around Chirchik, while tiger transgressioas in Ferghana valley were reported 
even in 1903. By 1910 to 1915 below Kyzyl-Orda tigers were still quite 
common, but north of it encountered rarely and irregularly. 

Commencing from the first and especially the second decade of the 
present century, tigers disappeared rapidly even in the lower reaches of 
Syr-Darya; the last tiger there was killed in 1933 (Kazalinsk region). Some 
stray animals were observed even up to 1937. After an interval of several 
years, tigers appeared for the last time in the lower reaches of the Syr- 
Darya at the end of 1945, when they were observed in a section from the 
estuary to Solotyub (slightly above Kyzyl-Orda). These animals had come 
from the Amu-Darya together with wild boar, which in that year migrated 
in very large numbers from the Amu-Darya delta along the shore of Aral 
Sea to the Syr-Darya. In the middle of the winter the animals disappeared 
from the Syr-Darya. The arrival of tigers from the Amu-Darya was again 
recorded in 1924. Tigers intruded far north on the Turgai River from the 
estuary of the Syr-Darya and the shore of the Aral Sea. 

Farther east tigers were encountered along Lake Telikul'sk and in the 
lower reaches of the Sarysu, and lived along the Chu from the mountains 
to its lower reaches; the animals lived there permanently and were not 
uncommon even early in the present century. The last two tigers on the Chu 
River were killed in 1912 but resident tigers were detected even in 1929 


(two pairs), 1936, and 1937. By 1940 they had disappeared (died). The 
tigers of the Chu River and the lower reaches of the Sarysu were linked 
with those of the Syr-Darya. At the beginning of the second half of the 
last century tigers were met with in the lower Talas and in the Biilikul' 
lake system, in the northern foothills of the Karatau and Kirgiz ranges, and 
also evidently on the slopes of Karatau.-'''* 
101 In the east tigers lived all along the Hi River up to the State boundary 

and throughout the system of creeks and channels in its lower reaches and 
delta, along the Karatal, Aksu, and Lepse, all along the reed jungles on 
the southern shore of Balkhash, along the Ayaguz (lower reaches), and in 
the rushes at Alakul. Tigers were mostly seen on the Hi, especially in the 
lower reaches; they lived along other rivers mainly in their lower reaches 
and estuaries. Even at the begiiming of the present century tigers appeared 
only sporadically in the lower reaches of the Karatal but litters were seen 
even in 1929 and the last encounter (catch) occurred in 1931. On the Aksu 
tigers were noted in 1909, on the Lepse still in 1930, and on the Ayaguz 
a litter was detected in 1908. On the Hi River tigers were first exterminated 
in the middle course (Iliisk). In the 1890's they were quite common there 
but appeared sporadically in the present century, although transgressions 
were reported even in 1929 and 1930. Tigers survived the longest in the 
lower reaches of Hi, living there until the 1930's. There were 10 to 12 of 
them in 1935 and 5 or 6 in 1939. Tigers had entirely disappeared (died) 
on the Hi River by 1948. Evidently, even transgressions from Kuldja are 
hardly possible nov*^. 

In the last century, even in the 1890's tigers were rarely sighted in 
the hills south of the region under description. The animals were reported 
in the Trans-Hi Alatau, the Ghu valley, the Kirgiz (Aleksandrov) range, 
and in particular on the southem slope to the Talasu in the Talas city region 
and in the Talassk valley, at Issyk -КиГ, in the upper reaches of the Naryn 
and on the Narynkol.-^^ 

The eastern edge of the Middle Asian region of tiger habitation is formed 
by the Tarbagatai [range], Zaisannor [Lake] and the lower reaches of the 
Chemyi Irtysh, and the Kurchum valley. Here animals were encountered 
only in the last century, becoming rare even toward the end of it.-^^ 

-^■^ According to data of Rychkov. 1762; N. Severtsov, 1873; Hem, 1891; Zamdnyi, 
1915; Sokolov, 1924; G.V. NikoFskii, 1930; Ognev, 1935; Sludskii, 1939, 1950, 1953, 1953a, 
and 1966; Kuznetsov, 1948; and others. 

^^According to N. Severtsov (1873) the occuirence of tigers there was common and 
permanent. In his view they were encountered only in summer in the alpine zone. 

^^According to data of Brandt, 1856; P.P. Semenov, 1865; N. Severtsov, 1873; Finsch 
and Brehm, 1880*; A. Nikol'skii, 1887; Alferaki, 1891; Slovtsov, 1897; Shostak, 1927; 
Shnitnikov, 1936; D. Demenfev, 1938; Sludskii, 1939, 1953, 1953a, and 1966; Kuznetsov, 
1948; P.I. Semenov-Tyan-Shanskii, 1948; and others. *Not in Literature Cited— Sci. Ed. 


Tigers are capable of negotiating very long distances and have been 
seen as intruders far from the region of their normal habitation. Often 
they find themselves in places unfavorable for their survival, on terri- 
tories with an environment altogether alien to them (open steppes and 
deserts). In Middle Asia and Kazakhstan many distant intrusions of tigers 
are known. Thus, along the northern edge of the Kyzylkum and the eastern 
bank of the Aral Sea they crossed time and again into the lower Syr-Darya 
(evidently also vice versa), which is about 500 km as the crow flies. In 
1945 it was conclusively established that tigers coming from the Amu- 
Darya to the Syr-Darya following wild boar traveled about 1,000 km in 
about two to three months. 

In the west transgressions are known in Turkmenia in the Great Balkhan 
(1936) and even the northern shore of the ICara-Bogaz-Gol bay (Shev- 
chenkovsk region, Guryev district; tigress and two cubs in 1947). In the 
Great Balkhan tigers undoubtedly traveled from Iran, to Kara-Bogaz-Gol, 
evidently along the southern Ustyurt chink [arroyo] from the Amu-Darya 
deha. The animals in both instances would have covered great expanses 
of desert. Transgressions were known from the Amu-Darya delta along the 
southern Ustyurt chink, and in the Ustyurt in the region of the road in 
Kungrad. Tigers have time and again occurred even on the southeast coast 
of the Aral Sea in the northern Kyzylkum and at Kuvan-Darya. From the 
Amu-Darya they evidently extended even to the Ural [river] estuary.**^ 

In Kazakhstan and adjacent parts of western Siberia tigers intruded 
at several places evidently from Pri-Balkhash, probably from the Chu and 
Sarysu Rivers, and from the Zaisan region. In the southern Altai and Pri- 
Altai regions tigers were reported at Altai (Katon-Karagai) station, 
Bukhtarminsk, Ust-Kamenogorsk, Semipalatinsk, Zmeinogorsk, Loktevsk 
Zavod on the Alee River, at Biisk (52°30') and even Barnaul (53°), and 
at other places in this same region (Setovka, B. Biisk u.; Pokrovskii Zavod). 
102 Along the Irtysh the animals reached Pavlodar and Yamyshev (about 50 
km above Pavlodar). These intrusions were reported in the last century, 
mainly in the early part; transgressions in the southern regions were not 
very rare. 

West of Irtysh, intrusions have been recorded for the region north of 
Balkhash, the Karkaralinsk mountains, toward Akmolinsk, and even into 
the Kokchetav mountains (Sedel'nikov and Borodin, 1903; Sludskii, 1953). 

•^^"Close to the estuary he (Yaik) was overtopped by tall reeds, where wild boar and 
tiger found refuge" (A.S. Pushkin, //wto?^' ofPugachev, vol. I). This lone reference of Pushkin 
has no reliable confirmation in zoological literature. There is, however, a reference to the 
habitation of tigers in the region of nomadic Little Order Kazakhs (Brandt, 1856). These 
animals were sighted within the boundaries of the UraFsk district, and in part in the Turgaisk 
and Syr-Daiyask districts, and in Orenburg and Astrakhan provinces. 


The last of these sites is about 700 km north of Balkhash. It is possible, 
however, that at least some of these intrusions were associated not with 
the habitat of the animal in the Balkhash region but with the Chu region, 
i.e., Sarysu and Syr-Darya. Such distant settlements were, in the last century, 
facilitated to some extent by a much broader past distribution of wild boar 
in the north (see Volume I of this monograph). 

In eastern Siberia (Fig. 50) tigers nowhere settled permanently or, if 
they did, it was so long ago that there exists no direct evidence. Tiger 
occurrences in Trans-Baikal and Cis-Baikal (see below) were, and continue 
to be, mere transgressions. There is no positive information about intrusions 
in the early part of the last century into the headwaters region of the Yenisey 
(Tuva, "Uryankhaisk Land"; Radde, 1862). 

102 Fig. 50. Distribution of tigers in eastern Siberia and along the upper Amur. 

1 — places and years of occurrence; 2 — localities in Pri-Amur (see Fig. 53); 

3 — references to Verkhoyansk mountain range and unconfirmed information about 

the upper reaches of the Irkuta, Oka, and Tuva. V.G. Heptner. 


Far Eastern distributional region. This area lies in the Pri-Amur and 
Ussuri regions; the Trans-Baikal region is zoogeographically identical to 
this one (Figs. 51, 52 and 53). All these regions of the range are linked 
with northeast China (Manchuria). 

The northern boundary of the habitat of the animal in this area in the 
historic period can only be tentatively established. Adequate information 
does not exist (available only from the commencement of Russian 
investigations, i.e., in the second half of the last century). Furthermore, 
multiannual fluctuations of the boundary inevitably resuhed from frequent 
transgressioas beyond the area of permanent habitation, natural causes, and 
103 disturbance by man. It is also significant that tigers lived and continue to 
live in the Far East not in isolated pockets along rivers in the desert as 
in Middle Asia, but over a large expanse with relatively uniform forest 

In the west, the reconstructed northem boundary of permanent habitation 
of the tiger commences on the Amur at the mouth of the Khumaerkhe 
(Kumara), flowing into Amur from the right. It runs east and southeast, 
encompassing the lower reaches of the Zeya and Bureya, its left tributary 
the Tyrma, the southern part of the Bureinsk range, the Bidzhan and Bir 
basins, the lower and middle reaches of the Urma (on the north a tributary 
of Urma, the Berendzha) and the Kur', and emerges at the Gorin. At the 
mouth of this river or slightly below it, the boundary crossed the Amur 
and, avoiding the Khungara basin to the south, extended toward the Tatarskii 
Strait. It is highly probable that the boundary actually crossed, at least 
at some places, slightly farther north, probably along the upper reaches 
of the Amguna, dov^nstream along the Amur (at 52° in the region of Lake 
Kizi), and so on. South of this line tigers occurred everywhere right up 
to the Korean border, although absent at places in the high mountains of 
the Sikhote-Alin. 

This reconstructed range corresponds roughly to the situation obtaining 
in the middle of the last century. As a result mainly of direct persecution, 
and in part felling of forests and extermination of wild ungulates, especially 
wild boar, the range was greatly ahered. On the whole, from the end of 
the last and the beginning of the present century, there has been a significant 
reduction of the range, mainly in the north. Thus, by 1940 the northern 
boundary of permanent habitation in the Ussuri region extended in general 
along the Iman, and the animal was almost absent throughout the northern 
part of the Ussuri region, along the Khor and Bikin. This reconstructed range 
became the region of later occurrences (but not all); as a result of reduced 
persecution during the war, establishment of preserves, and enforcement 
of measures for their conservation, the range of tiger enlarged somewhat. 
By 1950 tigers were found in the north along the Bikin and the southern 


104 Fig. 51. Change in some range boundaries of tiger in the Far East. 

1 — northern boundary of permanent habitat; in part of permanent and regular 
intrusions in the first half of the nineteenth century'; 2 — approximate northern 
boundary in 1940, tlie period of utmost recession; 3 — boundary in 1950; 4 — some 
peripheral points of intmsions from 1920 to 1962; 5 — from 1952 to 1962 
(according to data of N.V. Rakov); 6 — points of earlier intrusions; 7 — occurrences 
in Sakhalin (nineteenth century, no exact site). V.G. Heptner. 

tributaries of the Khor and along the coast reached Samarga where the animal 
had long been absent (Fig. 51). 

After 1950 enlargement of the range of tiger conrinued and by 1969 
(data of S.P. Kucherenko and A.G. Pankrat'ev for December, 1968) 
comprised three sections (Fig. 52). Maximum enlargement occurred in the 
Sikhote-Alin. The region of permanent habitation extended there roughly 

105 Fig. 52. Distribution, nature of occurrence, and comparative numbers of tigers 

in Pri-Amur and Ussuri regions at the end of 1968. 

1 — region of permanent habitat; 2 — sections of maximum population density; 

3 — region of regular temporary occurrence; 4 — points of distant occurrence in 

the 1960's; 5 — direction of more or less regular migrations; 6 — route of particular 

distant occurrence. S.P. Kucherenko and A.G. Pankrat'ev. 


up to 46° N. lat. (the upper Kema); one extension ran northward on the 
eastern slope of the mountain range along the coast, extending slightly north 
of the mouth of the Samarga (about 47°30'), and another ran along the 
western part of the Sikhote-Alin north of the latitude of Khabarovsk (almost 
to 49° N. lat.). 

This range was surrounded by a region of temporary but regular trans- 
gressions. In the west this ran initially in a narrow strip along the western 
slopes of the Sikhote-Alin and along the Ussuri, and they continued as a 
broad projection to Komsomol 'sk-on- Amur. In the middle of the Sikhote- 
Alin this region extended approximately to 48°20' N. lat. and along the 
eastern slopes of the range, to 48°45', i.e., almost to Sovetsk Gavan'. 
Northward from there some individual irregular intrusions extended in the 
1960's up to the upper Tumuin, Amur at 51° N. lat., the sources and 
midcourse of the Gorin (Goryun; left tributary of the Amur, entering below 
Komsomol'sk), and the Duka River the source of the Amguna in general 
slightly north of 51° N. lat. The region of maximum density of the tiger 
population occupied almost the whole of the southern part of the region 
of permanent habitation, roughly from the upper Armu and upper Kolumba 
at the source of the Iman (slightly south of 46° N. lat.). 

A very small separate section of tiger habitat extended in the form of 
a narrow strip, along the southernmost section of the state boundary from 
the level of Lake Khanka to the latitude of Vladivostok or slightly farther 
south. There are two regions of permanent habitation here and two of regular 
intrusions from the west. This section was isolated at the end of the 1960's 
from the main Sikhote-Alin range described above. 

The third section of tiger habitat lies inside the large southern bend 
of the Amur. Commencing at the Amur, southwest of the town of Obluch'e, 
104 it extends initially east almost to the town of Birobidzhan and later southeast, 
mnning again toward the Amur west of the mouth of Bidzhan. The range 
thus encompasses the southeastern part of Bureinsk range and Pri-Amur 
lowland to the east. The central part of the section adjoining the Amur 
represents the region of permanent habitation; it is surrounded by the region 
of regular intrusions. Some more distant intrusions from this part of the 
range extended in the 1960's to the central parts of Urma basin and the 
midcourse of the Kur'.-""^ 

As in the earlier period, in the 1960's also there were regular migrations 
of tiger between northeast China (Manchuria) and the Soviet Union. At the 
very end of this decade such migrations took place in the Amur bend, the 

^^'By 1970 the situation in this pait of the range and on the right bank of the Amur 
had undergone a change. This region, which invariably received migrant animals coming 
from beyond the Amur, is evidently devoid of tiger or is becoming so. In any case, in the 
winter of 1968 to 1969 not a single track was detected in Birobidzhan (A.G. Pankrat'ev). 


region of the mouth of the Bikin and Ussuri, and in the section of the range 
in the extreme southeastern Ussuri region adjoining [lake] Khanka (S.P. 
Kucherenko and A.G. Pankrat'ev). 

The enlargement of the range described above from the time of its utmost 
reduction in 1940 as well as the increase in tiger population, which took 
place in spite of a rapid increase in human settlements and intense cultivation 

105 of the country (mainly extensive exploitation of forests), are attributed 
primarily to some of the conservation measures implemented after the war. 
In spite of improved natural conditions, which for a long time were not 
present in the west, tigers even here would have faced rapid extinction had 
such measures not been implemented. By the end of the 1960's a fairly 
extensive territory was occupied by settled breeding populations and a fairly 
large number of regular intmsions. They are such that through very careful 
regulations it is possible to conserve the animals. 

106 Tiger intrusions into the Far East always were very extensive, reaching 
far beyond the limits of the reconstructed boundaries described above, 
especially in the past (Figs. 50 and 53). However, in our century and even 
in the past decade, tigers penetrated quite far, especially when one recalls 
that the northern boundary of the region of permanent habitation within 
the Soviet Union had receded southward markedly (see above). These tigers 
have been sighted in different years somewhat above the mouth of the 
Kumara on the Amur, at Selemdzha around 52°, north of the Berendzha 
River, a tributary of Urma (about 49°30' to 50°00'; probably this is not 
an intmsion but a rare, more or less permanent habitation); at Niman on 
the right tributary of the upper Bureya (around 51°30'), on the upper Amguna 
(slightly south of 51°), on the Amur below the mouth of the Gorin, at Lakes 
Kadi and Kizi on the lower Amur, and in certain other places, extending 
sometimes to 54°, i.e., up to the latitude of the Amur estuary. 

From 1920 to 1962 tigers transgressed into an extensive region north 
of the Amur, as may be tentatively judged by the following reports: the 
region of Yerofey Pavlovich and Solov'evsk; slightly west of the town of 
Zeya on the Zeya; between the upper Zeya and its right tributary the 
Mul'shuga; the left bank of the upper Zeya at the meridian of the town 
of Bomnak; slightly east (on 130° E. long.) on the same latitude; the 
headwaters of the Nora, a right tributary of the Selemdzha; the headwaters 
of the Selemdzha; the right bank of the midcourse of the Amguna on the 
meridian of L<ike Chukchagir (left bank of the Amguna); left and right banks 
of the Amur slightly below the mouth of the Gorin; the upper Tumnin (data 
of N.V. Rakov; Fig. 51). 

107 Intmsions in the 1960 's (to 1968 inclusively) for the same region were: 
midcourses of the Urma and Kur', the Duka River in the headwaters of the 
Amguna, headwaters and midcourse of the Gorin, right bank of the Amur 


106 Fig. 53. Distribution of the Amur subspecies of tiger, Panthera tigris altaica Temm. 

1 — range boundaries at end of the nineteenth century; 2 — range boundaries by 

the 1930's; 3- 

-sections outside the range; 4 — separate long dispersals of tigers. 
K.G. and V.K. Abramov. 

below Komsomol'sk, and headwaters of the Tumnin (S.P. Kucherenko and 
A.G. Pankrat'ev; Fig. 52). 

Some intrusions, partly in the remote past and partly in our day, extended 
significantly farther. In 1944 tiger intrusions were reported on the upper 
Uchur (right tributary of the Aldan) in southeastern Yakutia, approximately 
57°20' (D. Ivanov), and on the Amga River (right tributary of the Aldan) 
at 60° N. Lat. Evidently the report of the sighting of a snow leopard on 
the Olekma 300 km from its mouth (Pallas, 1811; see chapter on snow 
leopard) also refers to a tiger. An exceptionally distant intrusion was reported 


in 1905 when two tigers were encountered on the Aldan 60 and 80 km 
below Ust'-Maya (about 60°45'). Less reliable is information about the 
appearance of tiger in the Verkhoyansk range, evidently south of it. Tiger 
transgressions are also known for Sakhalin.-*^^ References to normal 
occurrence on Sakhalin are erroneous, certainly (Fig. 50). 
108 Evidently, the majority of these intrusions were linked with northeastern 

China (Manchuria) and only a portion of the eastern points were reached 
from the southern bend of the Amur and the Ussuri region. 

The distribution of tigers in Trans-Baikaliya is quite distinctive. They 
occur here only as intruders, evidently from the Great Khingan [mountains] 
in northeast China, and although they sometimes stay for quite some time, 
they never breed. Intrusions extend into the easternmost (often) and southern 
parts of the Trans-Baikaliya.^° Reports are on record of tiger at the 
confluence of the Shilka and Argun, at other places along the Argun, i.e., 
from its mouth upstream to the Tsurukhaitui (about 50°15'), at the large 
Nerchinsk factory, on the Gazimur River near its confluence with the Argun 
(Kuchugai), at Gorbitsa on the lower course of the Shilka (about 53°05'), 
at Aksha on the upper Onon (near the State boundary — about 50°15'), and 
on the Ingoda near Uleta (about 100 km straight up the river from Chita). 

Intrusions into Trans-Baikaliya were not rare. Much of the information 
pertains, however, to the first half and middle of the last century. At the 
end of the nineteenth and the first half of the twentieth century, definite 
information about the appearance of tigers in Trans-Baikaliya is not available 
but without doubt intnisions did occur. However, in December, 1953 a tiger 
was killed at Kaktolga on the Gamimur (52°45' — slightly above Kuchugai), 
another on the bank of the Shilka in the Mogochinsk region, and a third 
on December 29, 1953 at Balei (south of Nerchinsk on the Unda River). 
In that year alone eight tigers were sighted in southeastern and eastern 

Trans-Baikaliya is evidently a region through which tiger used to 
penetrate even farther east- and northeastward. Thus, in 1828 a tiger was 
noted at Balagansk on the Angara north of Irkutsk, and tigers occurred twice 
in the thirties at Nizhne-Angarsk at the northern extremity of Baikal (56°30'; 
Sludskii, 1953a). It is possible that some of the deep intnisions mentioned 
above also occurred through Trans-Baikaliya (Fig, 50). 

-'^^he Far East part of the range is from Przhevalskii, 1870; Baikov, 1925 and 1927; 
Afanas'ev, 1934; Salmin, 1940; Kaplanov, 1948; Sludskii, 1953 and 1966; V.K. Abramov, 
1962 and other works; and data of K.G. Abramov, G.F. Bromlei, S.P. Kucherenko, A.G. 
Pankrat'ev, and V.G. Heptner. For other tiger intrusions see the section "Biology". 

^It is possible that on several occasions animals may have entered Trans-Baikaliya 
along the Amur valley but in general this is rarely tioie; concerning tiger habitat on the Amur 
in the expanse from the mouth of the Kumara upstream to the confluence of the Shilka with 
the Argun, there are no data. 


Geographic Range outside the Soviet Union 

The range outside the Soviet Union covers northern Iran, Afghanistan, much 
of India except the northwestern desert regions of Rajputana, Punjab, Kutch, 
and Sind; Sri Lanka, Indochina, Malacca, Sumatra, Java, and Bali; the Hi 
basin in Dzhungaria, Lobnor, Tarim and Yarkand-Darya in eastern Turkestan; 
northeast China in the west including the Great Khingan, and the eastern 
part of China. Tigers are now extinct over large areas (China, except the 
northeast and extreme south, much of India, Bali, etc.). If one considers 
range as a whole, then the tiger is, apparently, extinct over most of it.''^ 

From the zoogeographic point of view the tiger represents a typical 
south Asian form, most probably with southeast Asia as its place of origin. 
Its present-day (historic) range is the result of colonization by two distinct 
groups of animals; one group via the northeast through eastern Asia, and 
the other via the northwest through the Near East (Iran and Caucasus) and 
Middle Asia. Both these directions circumvented the Central Asian highlands 
and Tibet. Extension into Semirech'e, Altai, and Kashgaria on the one hand, 
and into Trans-Baikaliya, on the other, was a secondary phenomenon; these 
represent the terminal areas of two groups of animals coming from opposite 

According to some authors the tiger as a species is associated in origin 
and dispersal with eastern Trans-Baikaliya, northeastern China (Manchuria), 
and Pri-Amur (Baikov, 1925; K.G. Abramov, 1965). The present-day range 
is considered the result of colonization through eastern Asia southward and 
through southern Siberia westward to Turkestan and the Caucasus (Mazak, 
1965). This view is unacceptable not only from the zoogeographic viewpoint 
but also because it is based on an incorrect interpretation of the fossil 
109 remains of large Pleistocene cats of Siberia. They were long supposed tigers 
while, in fact, they were cave lions {Panthera spelea Gold.), which were 
widely distributed not only in the Palearctic but also in the Nearctic regions 
(N.K. Vereshchagin). Tiger remains are known only from the Upper Pleis- 
tocene and Holocene strata of the southern Ussuri region (Vereshchagin 
and Ovodov, 1968). In the Caucasus the tiger was evidently a more recent 
colonizer, appearing in the Holocene period (Vereshchagin, 1959). Fossil 
remains have not been recovered there; nor, in particular, have they been 
discovered in the Binagedinsk excavations. (V.G.) 

Geographic Variation 

Geographic variation of the tiger is not very sharp but nevertheless manifested. 

^^The reported occurrence of tiger in Sudan, with even a description of a special fonn, 
P. с siidanesis (Daraniyagala, 1952), lacks credibility and warrants no scientific discussion. 


107 Fig. 54. Reconstructed species range of the tiger, Panthera tigris L. Territory of 

permanent iiabitation and regular, relatively shortniistance intrusions. Dots indicate 
deep intnisions beyond the northern boundary of the range; crosses denote regions 
occupied or visited by tigers in the Middle Ages. V.G. Heptner. 

Until recently about nine subspecies were recognized, with six in continental 
Asia (Ellerman and Morrison-Scott, 1966). Usually three, sometimes even 
four subspecies are assigned to the Soviet Union. The number of names 
is around 20. Soviet zoologists have always recognized only a single form 
for the Caucasian-Turkestan part of the range. The separation of the Balkhash 
tiger into a distinct subspecies (trabata) has not been recognized at all. For 
the Far East two subspecies have long been recognized — the northern 
"Amur" {longipilis) and the more southem "Korean" (coreensis; Ognev, 
1935) even though in the 1920's it was demonstrated that these "subspecies" 


are merely individual variations and that only one form of tiger inhabits 
the Far East (Baikov, 1925). This was convincingly proved later (Stroganov, 
1962). Yet even now some researchers persist in recognizing the earlier 
two forms of Far East tigers (Bobrinskii, Kuznetsov and Kuzyakin, 1965). 
The total number of tiger subspecies is evidently eight, of which two are 
known in the Soviet Union. 

By and large, the geographic variation of tiger is quite regular. The 
largest form comes from the most northem parts of the range of the species 
(Amur), while the smallest inhabits the Malay Archipelago; tigers of 
Turkestan, Iran, India, southern China, and Indochina occupy an intermediate 
position in size and some other features (type and color of coat, and extent 
of difference between winter and summer coats). Bergmann's rule and iasular 
characteristics are well exhibited. 

The following forms inhabit the Soviet Union. 

1. Amur tiger, P. t. altaica Temminck, 1844 (syn. amurensis, coreensis, 
mikadoi, mandslmrica, and longipilis). 

Size large, exceeding on the average the size of the largest of Indian 
tigers. Build heavy and massive; anterior part of trunk and forelegs parti- 
cularly well developed. Head relatively heavy and large. ^^ 

The main background color of the winter coat is a relatively light, 
ocherous-yellow, with a fairly intense admixture of reddish-rust. In general, 
however, color is very variable and in extreme cases may be ocherous- 
yellow or fairly bright reddish-rust, especially on the back. In the summer 
coat this color is significantly brighter and more reddish. In general, the 
Amur tiger is one of the most variable forms of the species insofar as color 
is concerned. '^^ 

Underside of body and inner surface of legs white and white color on 
flanks usually extends quite far up. White fields around eyes quite large. 
Dark bands vary rather significantly in number, size, form, and partly in 
112 color. Usually, they are relatively narrow, long, often bifurcate at the ends, 
or occur in pairs throughout. Much shorter lenticular bands, pointed at the 
ends, with a gap in the middle and not reaching either the sacrum or the 
abdomen, are frequently seen. Number of stripes relatively small, i.e., they 
are not close-set (Figs. 55, 56 and 57). 

Stripes on body black but on thighs, base of tail, and sometimes even 

■^These features of general appearance do not lend themselves to an accurate 
description but are given by many taxonomists as characteristic of Siberian tiger. The long 
coat is partly responsible for the general impression. 

^-^For a long time local Russian huntere distinguished tvi'o "subspecies" of tiger — 
bright rust-colored and dull-colored. These 1л\'о types of coloration provided the basis for 
differentiating the Гл'о subspecies — "Korean" (coreensis) and '"Amur' ' (longipilis). It was 
thought that bright-colored animals were smaller and their yellow undeifur shorter. 

110 Fig. 55. Color variation in tlie Amur tiger, Panthera tigris altaica Temm. Typical 

pattern of broad, relatively sparse stripes. 

A— No. S 55281, Khabarovsk region, 1952/1953; B— No. S 55280, same; C— 

No. S 55279, same; D — No. S 2%54, Amur. All specimens from the Zoological 

Museum, Moscow University. Photographs by D.M. Vyazhlinskii. 

•Чт-гт/ч/щ^, I » 

I™ "■,■ 

Fig. 56. Color variation in the Amur tiger, Panthera tigris altaica Temm. Rarer 
specimens with a pattern of relatively close-set and narrow stripes. 

A— No. 3029, southern Ussuri region, Sidemi River, February 20, 1884; B— No. 
2980, central Amur; C— Amur-Ussuri region; D— S 50146, left tributary of central 
Amur (around 48°30' N. lat.), 1940. First two specimens from collection of the 
Zoological Institute, Academy of Sciences, USSR and other two from the Zoological 
Museum, Moscow Univeraty. Photographs by D.M. Vyazhlinskii and L.G. Turova. 


Fig. 57. Color variation in the Amur tiger, Panthera tigris altaica Temm. More 

or less intemiediate pattern between types shown in Figs. 55 and 56; relatively sparse 

bands in anterior part of body and narrow and close bands in posterior part. 

Left— No. S 55284, Khabarovsk region, 1952/1953; Right— No. S 29664, female, 
Kliar River, tributaiy of the Pikhtsy, ti'ibutaiy of the Amur (49° N. lat., about 76°30' 
E. long.), March, 1938. Collection of the Zoological Museum, Moscow University 

(see Fig. 45). 

flanks usually not pure black but different shades of brown or brownish. 
Head, middle of back, abdomen, and distal half of tail generally pure black. 
In animals with very vivid color, the pattern very shaф; in light-colored 
animals usually less 811аф, especially on flanks. Moreover, pattern less distinct 
in animals in long-haired winter coat. 

Rings on tail (not counting black tip) usually eight or nine; they are 
generally paired (with a light-colored transverse gap along the base). In the 
basal region toward the top usually (more so evidently in males) a pattern 
in the form of angles with the apex turned forward occurs (Fig. 58). 

The color of subadult tigers does not differ from that of adults; tigresses 
at the age of several months (three or four) are paler. 

Seasonal coat dimorphism is extremely sharp, and a very characteristic 
feature of the subspecies. Summer coat hair in length and density barely 
differs from that of southern tigers (India), but the long and dense, even 


113 Fig. 58. Predominant color type of fails in Turanian tiger, Patithera tigris virgata 

Matschie (left) and Amur tiger, P. t. altaica Temm., after K.K. Flerov and I.M. 
Gromov, 1934. Sketch by N.N. Kondakov. 

slightly shaggy winter coat imparts to the Amur tiger an altogether special 
appearance. In summer, males, though not all, possess slightly longer hair 
on the neck and throat, and whiskers are faintly perceptible. In winter, however, 
in the majority, mainly males, the mane is greatly elongated as is the hair 
throughout the neck; whiskers are also very long. The tail becomes very 
thick and its diameter may reach 10.0 to 12.5 cm (at the base). In the summer 
coat the hair on the back is 15 to 17 mm long, along the top of the neck 
113 30 to 50 mm, on the abdomen 25 to 35 mm, and on the tail 14 to 16 mm; 
length of whiskers 70 to 80 mm. In winter the hair on the back is 40 to 
50 mm long, along the top of the neck 70 to 110 mm, on the throat 70 to 
95 mm, on the chest 60 to 100 mm, on the abdomen 65 to 105 mm, and 
on the tail 35 to 50 mm; length of whiskers 90 to 1 15 mm (Mazak, 1967). 
The size of Amur tigers varies markedly. The range of variation among 
males is particularly wide; they are considerably larger in size and heavier 
in weight than females. It is also significant that tigers live long and continue 
to grow for a very long period, practically throughout their lives. Under 
favorable conditions the animals appear well fed and even obese. Maximum 
weight of the Amur tiger is 390 kg (Baikov, 1927) and 384 kg (male; V.P. 
Sysoev, 1952).'^ It is possible that these figures are somewhat exaggerated; 

^A weight of 400 kg has even been reported in an exceptional case (Baikov, 1927). 


in any case, they pertain to large old animals, which are rare at present. The 
existence of animals weighing 325, 340, 350, and 360 kg has, however, been 
established (Baikov, 1925; Bel'skii and Bromlei, 1953; Mazak, 1965, cited 
from Sludskii, 1966). Males weighing up to 320 kg and females up to 180 
kg are considered a fairly "normal" maximum (Baikov, 1925). This figure 
is probably high for females, however. Some individual animals, as pointed 
out, greatly exceed this maximum. This is particularly true of the past, when 
tigers were bigger and hved longer. Early in the present century, "a splendid 
male of moderate size, well fed, and even obese . . . probably attained 160* 
kg" (its total length was about 360 cm— "18 chetvert"; Baikov, 1927). 
The maximum known body length of Siberian tiger is 317 cm (along 
the curve of the back). With a tail length of about 100 cm, this giant measured 
a total length of slightly less than 420 cm (Barclay, 1915; cited from Mazak 
and Volf, 1967). A body length of 300 cm and a total length of 375 cm 
(i.e., body length about 275 cm [sic]; a tiger of "colossal measurements"; 
Baikov, 1927), have also been reported. Based on these data the "normal" 
maximum size of Amur tiger is as follows: body length (along curve) of 
males up to 290 cm and of females to 190 to 200 cm (probably slightly 
more); tail length to 1 15 cm; length of hind foot in males to 40 cm or slightly 
more and in females to 35 cm or somewhat more; transverse length of ears 
in males to 120 mm or more and in females about 100 mm; and height of 
males at shoulders to 115 or 116 cm, possibly even more.'^'' Data on several 
animals as measured by zoologists "in the flesh" are given in Table 1. 

'"Apparent eiTor in original Russian; sliould read 460? — Sci. Ed. 

^-''Height at the sliouldei^s is generally not stated by authors. An old male with the legs 
stretched (as suggested for measurements) reached 124 cm (Mazak, 1967). For the big tiger 
refen-ed to above with a body length of 317 cm, the height at the shoulders was given as 
140 cm (Mazak and Volf, 1967), which hardly seems likely. 

A critical analysis of some animals and data for specimens collected have been given 
by S.U. Stroganov (1962). The figures given in that work, published posthumously, are 
contradictoiy and partly enoneous (pp. 428 and 437, and others). 

The following are some additional published data on sizes of Amur tiger: weight of adult 
males 196 and 217 kg; male three-year-old, 115 kg (G.F. Bromlei); female three to four years 
old from central Amur (in Radde region), body length 178 cm and tail length 87 cm; male 
caught slightly downstream along the Amur, 180-185 cm and 93-94 cm (Radde, 1862); male 
of "colossal" dimensions (killed in 1911), length with tail 375 cm, height at shoulders 116 
cm, and weight of "cleaned" animal, 250 kg (Baikov, 1927); animal caught in the winter 
of 1927/1928 in Kirin province (northeast China), body length with tail 400 cm and weight 
325 kg (Baikov, 1929); female killed on December 16, 1927 near Modaoshi station on the 
Chinese-Eastern railway, body length 295 cm and weight 200 kg (usual weight of tigers 160 
to 200 kg; Baikov, 1928); male killed in 1953 in Trans-Baikal, body length 200 cm and weight 
195 kg; another from there 250 kg (Korneev, 1954); two tigers killed in Primor'e, body length 
with tail 325 and 310 cm; animal from around Ussuri, body length 284 cm, height at withers 



115 The skull of the Amur tiger is distinguished not only in its large overall 
size, but also in the great development of crests, especially the sagittal; 
the height and strength of the latter exceed those of other forms of tiger 
and the lion. Skull measurements are given in Tables 2 and 3.^^ 

1 16 Found in Ussuri and Amur regions and Trans-Baikaliya, and as a vagrant 
in Pri-Baikal and southern Yakutiya. 

Outside the Soviet Union, in the northern and eastem parts of northeast 
China (Manchuria), the Korean peninsula, and as a vagrant in the 
northeastern extremity of the Mongolian People's Republic. 

2. Turanian tiger, P. t. virgata Nliger {sic\ Illiger], 1815 (syn. 
septentrionalis, lecaqid [sic\ lecoqui], and trabatd). 

Compared with the Amur tiger, general build somewhat less massive 
and size, on the average, slightly less. 

Main background color of the skin variable but in general a brighter 
and more uniform reddish-rust, especially the summer coat, but evidently 
bright even in winter (Fig. 59). Dark stripes narrower, long, fuller, and closer 
set. Color of stripes a fairly distinct admixture of brown or cinnamon shades, 

117 sometimes even an "intense cinnamon" (Ognev, 1935).*^^ Pattern 
invariably pure black only on head, neck, middle of back, and at end 
of tail. Rings on tail usually arranged singly at least in terminal half; 
angular patterns at base of tail less developed than in the Amur tiger 
(see Fig. 58). 

Seasonal color dimorphism prominent but less shaф than in Amur tiger. 
Winter coat considerably lighter in color and paler than summer coat with 
a less distinct pattern. Hair on winter coat much longer and denser and 
coat may even be "shaggy". Length of hair on back in summer 8 to 13 
mm and on abdomen 20 to 30 mm; on nape, hair up to 20 to 50, and even 90 

(footnote 65 contcl) 

85 cm, and weight 221 kg (Antonov, 1950); another, body length 225 cm and weight 112 
kg; maximum dimensions of "Amur" tiger, body length 276 cm and tail 100 cm; "Korean" 
tiger, 300 cm and 80 cm; and weight of Amur and Korean tigers up to 340 kg (Baikov, 
1929; from data furnished by A.A. Sludskii). 

A large male caught in its fii-st year in the wild and raised in a zoological garden (Prague) 
weighed about 250 to 260 kg (Mazak, 1967). A group of three animals in a zoological garden 
weighed: 11-year-old male, about 317 kg; 8-year-old female, 286 kg; and 3-year-old male, 
about 299 kg (Mazak, 1967). These figures are erroneous, at least in the case of the female 
and young male, and constitute exaggerations. 

^^he sizes of almost all the skulls preserved in Soviet museums are given in the 
Tables; measurements which fall slightly beyond the indicated limits are not included. A 
skull (presei-ved in Harbin) has a maximum length of 400 mm and a zygomatic width of 
280 mm (Baikov, 1925). 

^^ Animal in which all the stripes are cinnamon-colored known from Ferghana. 



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Greatest length 

Condylobasal length 

Basilar length^ 

Rostral width^ 

Zygomatic width 

Interorbital width 

Postorbital width 

Mastoid width 

Length of lower jaw^ 

Length of upper camassial tooth 

Length of upper tooth row'* 





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115 Table 3. Skull measurements of female Amur tigers (from V. Mazak, 1967) 

Old adults^ 





Greatest length 




Condylobasal length 




Basilar length 




Rostral width 




Zygomatic width 




Interorbital width 




Postorbital width 




Mastoid width 




Length of lower jaw 




Length of upper camassial tooth 




Length of upper tooth row 




Note: Materials and measurements the same as shown in Table 2. 

4n V. Mazak's Table 1 (pp. 554 and 555) the animals are shown as old; in Table 3 
(p. 558) as adults. The former has been used here since it would appear to be more correct, 
judging from skull measurements (V.H.). 

mm long. In winter, hair on back may reach 30 mm or more in length/'^ 
In this subspecies hair evidently tends to grow much longer along the top 
of the neck (mane) and on the cheeks (whiskers). 

Data on the size and weight of Turkestan tiger are scanty. Maximum 
body length of males exceeds 200 cm, reaching 213, 224, and 217 cm, with 
a corresponding total length of 312, 324, and 309 cm, perhaps even more 
(see below; a body length of 270 cm has been recorded but probably was 
284 cm measured as a skin). Females smaller in size; however apparently 
their body length may reach 200 cm (total length, 267 cm) but normally 
ranges from 160 to 180 cm (total length 240 to 260 cm). The maximum 
known weight exceeds 240 kg but evidently could be greater. Thus, although 
the Turkestan tiger is generally smaller than the Amur, it nonetheless attains 
large proportions. 

The following information is available on the dimensions of some 

^^Conceming Trans-Caucasian tigers, K.A. Satunin (1915) reports: "All winter skins 
examined by me had very long and dense fur of dirty-yellow-light-brown color with indistinct 
dark stripes." "In winter (the fur) is very long and shaggy". "The very longest fur . . . 
was that of a huge tiger killed at the end of February 1899 in Prishibinsk section (Lenkoran; 
V.H.). Its fur was very dense and long but coarse and had some indefinite dirty-yellow-gray 
or brown color with a barely perceptible dark pattern; a reddish shade was not discernible 
at all . . . [I saw no] other animal with such long fur of a dull color". "... The shorter 
(the fur), the more vivid the reddish shade and the зЬафег the black stripes." 

116 Fig. 59. Variation in color of Turanian tiger, Panthera tigris virgata Matschie. 

1 — No. 3027. "Kirgiz steppe" (probably Semirech'e— V.H.); 2— No. S 33151, 
lower Hi, about 250 km above mouth, November, 1939; 3 — No. 2983, Lobnor 
(eastern Turkestan — V.H.), November, 1889; 4 — No. 8679, Ferghana; 5 — No. 
2981, "Turkestan,"" 1872; 6— No. S 52871; upper Pyandzh,Tadzhikistan, March 
30, 1950. Nos. 1, 3, 4, and 5 — pelts from collection of the Zoological Institute, 
Academy of Sciences, USSR; Nos. 2 and 6 — from collection of the Zoological 
Museum, Moscow University. Photographs by D.M. Vyazhlinskii and L.G. Turova. 


individual animals. Body length 162 cm, tail length 86.5 cm, total length 
248.5 cm (Altai, on Aleya River; Spasskii, 1820); male — body length 181 
cm, tail length 98 cm, total length 269 cm, weight 181 kg (southern Balkhash, 
1930; A.A. Sludskii); female — body length 250 cm, tail length 67.5 cm, 
total length 267.5 cm (Kirgiz range; Chekal'shchik); body length 172 cm, 
tail length 91 cm, total length 263 cm, weight 132 kg (Biisk district, 1839); 
male — length from ears (!) to base of tail 194 cm and tail length 89 cm 
(actual body length was evidently about 220 cm, total length about 309 cm; 
V.H.) (Syr-Darya around Karmakchaya, G-ii, 1887)^^; body length more 
than 284 cm^^ weight more than 240 kg (lower Hi, 1891, T.V., 1910); total 
length about 306 to 340 cm ("17 chetvert"; Kerki in the upper Amu-Darya; 
Gavrilov, 1896); total length 270 to 300 cm ("15 chetvert"; mouth of Amu- 
Darya; Kosarev, 1899) body length 217 cm (Kungrad, Amu-Darya delta; 
Kuznetsov, 1896)^^; body length 224 cm, tail length 100 cm, total length 
324 cm (Ferghana, 1903; Berg and Ivanova-Berg, 1951); female — total length 
259 cm (Pocock, 1929; cited by Mazak, 1965); adult male killed on January 
10, 1954 at Kaine-ICasyr on the Sumbar (southwest Kopet-Dag) — body length 
225 cm, tail length 105 cm (Shukurov, 1958); adult female — body length 
165 cm, tail length 95 cm, total length 260 cm, weight 135 kg (upper Pyandzh, 
Tadzhikistan; March 30, 1950); adult female — body length 165 cm, tail length 
76.5 cm, total length 241.5 cm, length of ears 11.0 cm, length of hind foot 
31.5 cm, weight 97 kg (Kyzylsu River, Tadzhikistan; June 26, 1950; 
Chemyshev, 1958); small tigress ("Theresa") acquired for Moscow 
Zoological Garden in 1924, a subadult from Iran which died in 1942 — body 
length 160 cm, tail length 86 cm, total length 246 cm, length of ears 11 
cm, length of hind foot 32 cm, weight 85 kg (Zoological Museum, Moscow 
University); male^ — body length 187 cm, tail length 90 cm, total length 277 
cm (Lenkoran'; skin measurements); female — body length 178 cm 
(Lenkoran'; skin); female — body length 157.5 cm, height at shoulders 
78 cm (Lenkoran'; Dinnik, 1914); body length 173 cm, tail length 110 cm, 
total length 283 cm (Trans-Caucasus: mounted skin); body length 161 cm, 
tail length 93 cm, total length 254 cm (Trans-Caucasus; mounted skin); 
male — body length 270 cm, tail length 90 cm, total length 360 cm ("huge 
tiger" from around Prishib, Lenkoran'; Febmary, 1899; skin just removed), 
Satunin, 1915.^2 

Maximum skull length in males 297.0 to 365.8 (385.0— V.H.) and 
female— 195.7 to 255.5 mm; condylobasal length in males 259.0 to 307.9 

^^Material gathered and furnished by A.A. Sludskii. 
^•^Probably measured from skin or total length (V.H.). 
^^ After Ivanova-Berg (1927). 

^■^his account about exhausts the information available on measurements of tiger of 
this subspecies, nowadays almost extinct. 


and females 225.0 to 263.2 mm; zygomatic width in males 219.0 to 254.0 
and females 183.0 to 203.2 mm (Mazak, 1965). 

As mentioned above, the dimensions of the Turkestan tiger are 
somewhat less than those of the Amur tiger, but this is true more with respect 
to mean dimensions of the general population. Large animals conforming 
to the above-mentioned maximum proportions of Amur tiger are not known 
in the population of the Turkestan form and are evidently absent. However, 
very large individuals are also found among the latter. Such for example 
is the Prishibinsk tiger listed above with a body length of about 270 cm.'^^ 
A tiger, killed on the Sumbar in Kopet-Dag on January 10, 1954 (stuffed 
skin in Ashkhabad Museum) has a body length of 225 cm and a greatest 
skull length of 385 mm; these measurements are considerably more than 
the known maximum for this subspecies and even slightly exceed the 
maximum for the Amur subspecies (see Table 2). The condylobasal length 
of this skull is 305 mm and zygomatic width 205 mm. The canine along 
the front curvature is 70 mm long, which is the maximum for RiLSsian tigers 
(V.G. Heptner). The skull of the animal from Lenkoran' (Belyasuvar) has 
a maximum length of 362 mm, zygomatic width of 248 mm, and interorbital 
width of 78 mm (Satunin, 1905). 

This form is found in Middle Asia and southern Kazakhstan, in the east 
(as vagrants) to the westem foothills of the Altai and eastern Traas-Caucasus. 
By the end of 1950's it had become extinct in Soviet territory and only 
some stray animals occur irregularly as intruders in the south-eastern Trans- 
Caucasus, southern Turkmenia, and southern Tadzhikistan (Amu-Darya). 

Outside the Soviet Union it occurs in the northern parts of Iran and 
Afghanistan, western Dzhungaria, Tarim Basin, and Lobnor (Kashgariya). 

The Turkestan tiger is a well-defined form which differs from the Amur 
tiger in several features. At the same time it has more features in common 
with the Indian tiger (P. t. tigris), evidently, than with the Amur tiger. This 
is reflected in general dimensions and in the nature of the coat. The summer 
coat in density and length of hair is similar to that of Indian animals, while 
the winter coat is longer and denser. The general shade of the coat is also 
similar but the stripes of the Turkestan subspecies are usually narrower, 
longer, and closer set. The color of the stripes can hardly be called pure 
black, as cinnamon or brownish tones predominate. 

Outside the Soviet Union, the following subspecies are recognized: 1) 
Chinese tiger, P. t. amoyensis Hilzh., 1905 — central and southern parts of 

^-^K.A. Satunin (1905) reports that it was "a tiger of immense proportions ... I saw 
it in the flesh. It appeared to me to be no smaller than the common Tuzemna horse". 


eastern China, except the extreme south; 2) Indochinese tiger, P. t. corbetti 
Mazak, 1968 — extreme southeast of China (Hunan, Kwangsi, Kwangtung 
provinces), Vietnam, lower Burma, Malacca, and evidently Cambodia, Laos, 
Thailand, and northeastern parts of upper Burma^"^; 3) Indian tiger, P. t. 
tigris Linn., 1758 — India, Assam; limits and relationship to preceding form 
not known; 4) Sumatran tiger, P. t. sumatrae Pocock, 1929 — Sumatra; 5) 
Javan tiger, P. t. sondaica Temm., 1845 — Java; and 6) Balinese tiger, P. 
t. balica Schwarz, 1912,— Bali.^-'^ (V.H.). 


Population. In the extreme northwestern part of the range and in Trans- 
Caucasus in the early eighteenth century tigers were still found in small 
numbers on the east coast of the Black Sea in the Kolkhid lowland and in 
119 Adzharia. They became extinct there in the first half of the nineteenth 
century. In the last century and early in the present one, tigers were seen 
from time to time in eastern Georgia — the last animal was killed in 1922, 
and in Armenia where one was caught in 1948 (Sludskii, 1966). In the middle 
of the nineteenth century in extreme southern Azerbaidzhan — in the 
Lenkoran' lowland and Talysh mountains, tigers still survived in significant 
numbers and also reproduced there. In the middle of the last century 10 
to 20 tigers were caught around Lenkoran' every year in autumn and winter 
(Brandt, 1856). In 1896 tigers were "not particularly rare" in Lenkoran' 
and Talysh regions and one or two animals were still being killed annually 
(Dinnik, 1914). 

In the present centur)' the tiger population in Trans-Caucasus decreased 
rapidly and the animals seen there only periodically as intruders from Iran. 
Only at Talysh do tigers still live more or less permanently. In Trans- 
Caucasus only two tigers were caught between 1920 and 1950 and about 
ten between 1950 and 1966 (Sludskii, 1966), including one in 1961 near 
Astara and two between 1963 and 1966 in the Lenkoran' region. In Talysh 
only single individuals have been encountered (mid-1960's; Lagidze, 1964; 
Aliev and Nasibov, 1966). 

In Turkmenia in the last centur}' tigers were fairly common in Tedzhen. 
Here they were encountered in the lowlands even in the 1880's and 1890's. 
Along the Murgab in the past they prowled in sufficient numbers but toward 
the end of the century were common only in the very headwaters and along 
the Kushka River. Tigers did not live in Kopet-Dag, but only entered there 
from Iran. They were frequently seen on the Atrek, Sumbar, and Chandyr 

^''The relationship of this fomi to neighboring ones, especially the Chinese fom;;. 
requires further confirmation. 


See "Geographic Range outside the Soviet Union", p. 129. 


119 Fig. 60. Habitat of tiger and wild boar in Semirech'e, old river bed of the Hi, 

Est-Bakanas River in the delta. Photograph by V. Marakov. 

in western Kopet-Dag. Along the Amu-Darya in the last century the animal 
was known everywhere; by the early twentieth century it was already a rare 
vagrant mainly in the upper reaches and extreme lowlands (V.G. Heptner). 

By the beginning of the present century in Turkmenia, tigers had become 
almost extinct except on the Amu-Darya. In the 1930's its population 
somewhat increased but not for long. From 1900 through 1968 only ten 
tigers were caught in all; of these, the last was killed in 1954 in western 
Kopet-Dag at Kaine-Kasyr (Heptner, 1956; Shukurov, 1958; Sludskii, 1966). 
The occurrence of tigers in Turkmenia is possible, and from farther in Iran 
and Afghanistan. 

In Uzbekistan tigers were comparatively rare even at the end of the 
last century. In the 1930's their number increased slightly; for example in 
the lower reaches of the Amu-Darya 12 to 15 animals were counted in 1942. 
120 Later, in spite of a ban on hunting, their population decreased rapidly. In 
1947 in the delta of this river, there were only five or six animals and the 
last was killed in 1950 or 1951.^^ 

^^See footnote no. 50 above, p. 116. 


120 Fig. 61. Reeds in places fomierly inliabited by tiger in Balkhash. Photograph by 

S.V. Marakov. 

In Tadzhikistan tigers were still common in the 1930's in the Pyandzh, 
Vakhsh, Kyzylsu, and other valleys, but are now extinct. In the early 1950's 
the number of tigers in Tadzhikistan would not have exceeded 10 to 15 
animals (Chemyshev, 1958), but probably Chemyshev erred and the number 
of animals far less. From 1942 through 1950 in Tadzhikistan only 10 animals 
were caught. In this republic tigers disappeared completely in 1953 or 1955. 


There is only one report, which needs confirmation, that the predator 
appeared in the Pyandzh valley in the Chubek region in 1963 (A.A. Sludskii). 
In Tadzhikistan tiger intmsions from Afghanistan are still possible. 

In Kirgizia, even in the last century, tigers were rare in the Issyk-Kul' 
region, in the Tei^sk Alatau, in the Chu valley, and along the Bol'shoi Kabin, 
and disappeared there roughly in the 1880's. South of this country they 
became extinct in the early 1890's (Sludskii, 1966). 

In the last century tigers were still common in Kazakhstan, in the Syr- 
Darya, Chu, Hi, Karatal, and other valleys, in Balkhash, Alakul; and Zaisan 
from where they intruded from time to time northward to Kokchetav, 
Pavlodar, Barnaul, and to the Ishim and Irtysh. In the Tien Shan tigers were 
regularly found, but were very rare in the southern Altai. In the last century 
and early part of the present tigers periodically occurred in Ustyurt, having 
entered there from the lower Amu-Darya. They were often encountered 
in the valley of the Syr-Darya. "They (tigers — A.S.) were found in large 
numbers in reeds around the Aral Sea and along the Syr-Darya and, occurring 
there together with humaas, perpetrated serious attack on camels and horses" 
(Rychkov, 1762). 

Already in the 1840's and 1850's the tiger population in Syr-Darya 
valley had begun to decline quickly, due to killing of the predators themselves 

. '^'> ^' 

121 Fig. 62. Reed thicket 4.0 to 5.0 m high in tiger habitat in Semiiech'e. Dzhel' 

Turanga, in the lower Hi. November, 1959. Photograph by A.A. Sludskii. 



and of wild boar, their main source of food, and destruction of the reeds. 
At that time "hunting teams", some officers and Cossacks from the local 
garrisons, killed 8 to 12 tigers in a brief period. In the early twentieth century 
in Syr-Darya this predator survived only at a few places. In five years (from 
1905) in the Syrdar'insk district only nine tigers were killed (Zarudnyi, 
1915) and their population continued to decline. The last two tigers were 
killed on the Syr-Darya, in 1924 in the Aral region, and in 1933 in the 
Kazalinsk region. Tiger tracks were, however, noticed even in 1937 and 
1945 when some animals entered into the lower reaches of the Syr-Darya 
from the Amu-Darya deha (Sludskii, 1966). 

In the Chu valley tigers were extinct by 1912 or at least 1916-1917. In 
the Hi delta, where tigers had survived for a particularly long period, only 
ten were found in 1935. The last was killed here in the winter of 1934/1935 
and the animal had disappeared totally from southern Pri-Balkhash in 1948. 
In the Zaisan depression tigers were already extinct in 1894. 

Thus tigers are exceptionally rare nowadays and seen only as vagrants 
in the Trans-Caucasus; they have disappeared completely from all the 
republics of Middle Asia and Kazakhstan. Intrusions are still possible into 
Tadzhikistan and extreme southwestern Turkmenia. 

121 Fig. 63. Upper Hi delta, habitat of tiger and wild boar. Roe deer were also very 

common in tliis region. Dzhel'-Turanga, Semirech'e. Photograph by V.S. Marakov. 


In eastern Siberia in the first half of the nineteenth century tigers were 
regularly seen at Baikal (Ehrenberg, 1831; Simashko, 1851; Sel'skii, 1856) 
and in southeastern Trans-Baikaliya, intruding from there far to the north, 
appearing from time to time even in the Verkhoyaask range (Silant'ev, 1919). 
Although not every year, one or two animals were caught in Trans-Baikaliya 
in some years, and tigers were caught from time to time even in the early 
twentieth century. Thus some were killed in 1912, 1926, and 1928. After 
a long interval they reappeared in Trans-Baikaliya, in 1953 and 1954. In 
1953 some eight animals were counted, of which two were caught (Sludskii, 
1966). From 1963 to 1966 tracks of the predator were found on the 
Uryumkan and Gazimur Rivers (Sinitsyn, 1966). Tigers penetrated into 
Trans-Baikaliya from the Great Khingan range of China. 

Farther east, in Amur basin, tigers were encountered in the last century 
and early part of the present one fairly regularly northward to 50° N. lat. 
They were common south of the Bureinskiy range on the Amur around 49° 
N. lat. (Radde, 1862). In the 1890's in the Ussuri region about 120 to 150 
tigers were caught annually (Silant'ev, 1898). Early in the present century 
the tiger population in the Far East declined rapidly, and by 1912, for 
example, only two animals were caught in the Amur region, and 55 in the 

122 Fig. 64. Tugais of tiuanga and tamarisk at Iman-baba, site of former tiger habitat 

in the Murgab valley. Turkmenia. April, 1962. Photograph by A.A. Sludskii. 



Primor'e (Baikov, 1927). The tiger population suffered a particularly great 
reduction in the Far East in 1929. 

Collectivization of agriculture and, later, measures undertaken for the 
conservation of tigers resulted in an increase in their population so that 
in the 1940's the animal was seen in regions whence it had disappeared 
long ago. Conditions for the reproduction of tigers remained favorable in 
the Far East until 1948; subsequently intense netting of cubs and shooting 
of adult animals decimated the population. Only 27 cubs were netted between 
1930 and 1938 in the region under description (Kaplanov, 1948), while 
56 were caught in the eight-year span between 1948 and 1956. From 1950 
through 1960 in the Primor'e region alone (southern half of the Ussuri) 
23 tigers were killed. 

The Game Inspectorate studied the tiger population in the Khabarovsk 
and Primor'e regions in 1956 and 1957. In the Khabarovsk region the area 
of tiger habitat was about 80,000 km^. The occurrence of 28 tigers was 
established over an area of 56,000 km-. In the Primor'e region 35 animals 
were counted in an area of 21,000 km^. 

In 1958 and 1959 in the Primor'e region an expedition of the Far East 

122 Fig. 65. Stieam and tugais in the Tigrovaya Balka preserve. Habitat of tiger, wild 

boar, Bukhara deer, and jungle cat. Lower Vakhsh, Tadzhikistan, October, 1960. 
Photograph by A A. Sludskii. 


Branch of the Academy of Sciences of the USSR, and the Central Game 
Board of the Russian Soviet Federated Sociahst RepubUc estimated the tiger 
population at 55 animals. These studies did not cover certain regions in 
which 8 to 10 or more animals lived. Evidently, in the whole of the Primor'e 
region 60 to 65 tigers lived in the first half of 1959. Their range covered 
23 administrative regions with a total area of 51,000 km^ (about 30% of 
the whole area of the region). In the Khabarovsk region (data of the Board 
of Game and Hunting) there were some 35 tigers in 1959 and a few more 
confined to the Amur basin. Thus in the first half of 1959 there were about 
100 tigers living in the Soviet Far East, constituting their total population 
for all of the USSR. 

The range of the tiger has undergone some changes in recent years in 
the Far East. As a result of a sharp reduction in population of wild boar, the 
predator has disappeared in the Amur region. In the Primor'e and a series 
of portions in the northern part of the tiger's range — in the upper Iman 
and Bikin, the population has declined somewhat. This reduction is explained, 
on the one hand, by the further human colonization of forest massifs along 
the Armu and Tatibe Rivers and the Iman basin, and on the other, by a great 

123 Fig. 66. Thickets of turanga, tamaristc, and plume grass, the site of tiger habitat 

in Tigrovaya Balka preserve. Lower Vakhsh, Tadzhikistan. October, 1960. 
Photograph by A.A. Sludskii. 


reduction in the population of wild boar in these regions due to inadequate 
yields from food crops and winters of intense snowfallfor several coasecutive 
years. However, in southern and southeastern regions of the Primor'e the 
tiger population has risen even in cultivated regions adjoining the Suchan, 
Maikhe, Tudagou, and other rivers; the animal is now encountered more 
frequently than in 1959. 

The total tiger population in the Soviet Far East, and thus for the Soviet 
Union as a whole, consisted of roughly 120 animals on January 1, 1965 
(Sludskii, 1966).^^ 

In the present centurj' the tiger population has decreased in countries 
adjoining the Soviet Union. Thus, in the regions of Turkey adjoining Georgia 
and Armenia several tigers were caught every year in the middle of the 
last century (Blit, 1863). At the present time tigers are extinct in Turkey. 
They have been preserved in northern Iran along the eastern slope of the 
Talysh and on the western banks of the Caspian [Sea], and they also inhabit 
Mazanderan, Gilan, Gorgan, and Khurasan provinces. In the 1930's in Iran 
80 to 100 tigers still survived but subsequently these numbers have declined. 
Tigers became ''quite rare" in the forests on the southern coasts of the 
Caspian. Apparently intense felling of forests has caused the animal to 
disappear altogether from Iran in recent years (Misonne, 1959; Ley, 1967). 

In Afghanistan, before the 1950's, tigers were common in the tugais 
along the left bank of the Pyandzh, from whence they frequently intmded 
into Tadzhikistan. In the last decade such intmsions have ceased, signifying, 
possibly, the disappearance of the animal from the former region. In the 
last century tigers were found in thickets along the Gerinid, Kunduz, and 
Murgab rivers, but today they have evidently disappeared there also. 

In China, Dzhungaria, and Kashgariya in the last century tigers were 
fairly common in the Tarim depression, along the Tarim itself, around 
Lob nor, and along the Khotan, Yarkand, Kyzylsu, Kashgar, Manas, Unmgu 
128 and other rivers, and in the Tien Shan. They had disappeared from the Tarim 
basin even in the 1920's and from the Tien Shan at about the same time. 
In 1959 tigers occurred with great rarity only along the Manas River, being 
absent already in other regions of Sinkiang; evidently at present they have 
now disappeared from there as well. 

Farther east, in Hailungjiang province (northeast China), adjoining the 
Soviet Union, and in the more southeriy Kirin province, there were still 200 

^^In early 1968 the total tiger population in the Far East was estimated at 122 to 148 
animals. The relatively poor increase in population, apart from other factors, is explained by 
illegal shooting in the remote regions. In 1966 to 1967 about 20 tigers were reportedly killed 
illegally (Kucherenko, 1970). Nevertheless the ban on hunting and enforcement of conservation 
measures have tripled the pre-1940 population level of the animal in the Far East (V.H.). 


123 Fig. 67. Turanga valley forest (tugai). Habitat of tiger. Lower Vakhsh. Tadzhikistan. 

Tigrovaya Balka preserve. April, 1966. Photograph by G.N. Sapozhnikov. 

to 250 tigers in 1958. As a result of intense felling of forests their population 
declined rapidly in this region. 

In the latter half of the last century about 150 tiger skins were exported 
annually from the Korean Peninsula. Today in the South Korea they are 
seen no more. In North Korea they have survived in regions adjoining Kirin 
province of the Chinese People's Republic; some 40 to 50 were supposedly 
still living there in the 1950's (Sludskii, 1966). 

At present world tiger reserves number 15,000 animals (Perry, 1964). 
Countrywise the distribution is as follows: Soviet Union — 120; Iran — 80 
to 100; possibly less; India and Pakistan — 3,000 to 4,000; Chinese People's 
Republic — 2,000; Korean People's Democratic Republic — 40 to 50; and 
Malaysia — 3,000. There is no information for other countries (Sludskii, 

Habitat. In the southeast Trans-Caucasus tigers were mostly confined 
to forests in Talysh lowlands in places where reed thickets along marine 
lagoons (mortsov) and streams were adjacent. These dense forests with 
abundant prickly vines could be negotiated only with the help of an axe 


124 Fig. 68. Thicket of giant plume grass {Erianthus purpurascens) at site of tiger 

habitat in "Tigrovaya Balka" preserve. Lower Vakhsh and Pyandzha. October, 
1960. Photograph by A.A. Sludskii. 

or scythe. Wild boar flourished in both forests and reeds but tigers pursuing 
them were encountered only rarely in the montane forests of Talysh. 

In Turkmenia, Uzbekistan and Tadzhikistan the main habitats of this 
predator are drainage basins of rivers and lakes, densely grown reeds attaining 
gigantic heights (5.0 to 6.0 m), and plume grass or tugai forests consisting 
of poplar, oleaster, and willow. All tugai trees are entwined with vines 
[Clematis]. In the lower stratum, cane, reed grass, licorice, camel's needles, 
and other plants flourish. These places abound in wild boar today, and 
formerly Bukhara deer were also common. At such places, where the 
approach to the river is across long expanses of solonchaks or sands, tigers 


126 Fig. 69. Imbishi area in Sudzukliin preseive — a common place for tiger residence. 

Nut pine-broadleaved taiga rich in Mancliurian wapiti and wild boar. Southern 
Ussuri tenitory. December, 1964. Photograph by V.E. Prisyazhnyuk. 

are absent or encountered only occasionally. Rarely these animals are seen 
in the foothills and the lower montane beh along valleys and gorges 
overgrown with pistachio, juniper, dog rose, etc. 

In Tadzhikistan tigers ascended to higher elevation from below 
extremely rarely. In some regions tigers lived fairly regularly close to human 
settlements. Relatively small reed thickets, covering several hundred hectares 
set between aids [villages] where wild boar roamed, were suitable for tiger. 
For example, in the Yakhsu River valley in Tadzhikistan, long inhabited 
by man with considerable land under pasturage, tigers lived permanently 
in the thickets in sections 1.0 to 3.0 km long and about 500 m wide; sections 
were located between rice and cotton fields. 

In the lower Syr-Darya and along rivers and large lakes in southern 
Pri-Balkhash, tigers lived in extensive reed floodplains alternating with lakes, 
streams, and series of sand knolls overgrown with saxaul, dzhuzgun 
[Calligofium], winterfat, and other plants, and also in the tugais. These areas 
abounded in wild boar and roe deer (in Syr-Darya, Bukhara deer replaced 
roe deer). 


126 Fig. 70. Montane taiga in the Tun'shi valley. Habitat of tiger. Sikhote-Alin preserve, 

Ussuri region. August, 1964. Photograph by V.E. Prisyazhnyuk. 

In Kazakhstan, and also in Kirgizia, tigers were sometimes encountered 
in montane belts, in summer ascending up to the permanent snow line. In 
the Kirgiz, Trans-Ili, and Dzhungarsk Alatau [ranges] they were caught 
in fir and juniper groves at heights of 2,500 to 3,000 m above sea level. 
In the mountains tigers were found even along valleys in thickets of tamarisk, 
willow, or sea buckthorn. These predators ascended the mountains in pursuit 
of wild boar and domesticated cattle which migrated there in summer. In 
Middle Asia and Kazakhstan habitats preferred by tiger roughly comply with 
the following requirements: 1) abundance of wild boar and Bukhara deer 
129 which serve as their main prey; 2) abundant available water, which is 
important for this predator that drinks several times a day; 3) availability 
of dense, impenetrable thickets in which tigers set up dens; and 4) low snow 
cover. These ecological requirements of tiger explain their absence in deserts 
far from water sources and only temporary or chance appearances in steppes 
and at high ahitudes in hills. 

Tigers live in the Far East under a different set of conditions. Here 
they are mostly confined to low mountains, especially in the last decade. 
In the last century they were even encountered at low altitudes but soon 
were displaced by man. Nowadays the most common habitats of this predator 
are montane river valleys and pad's overgrown with Manchurian-type 


127 Fig. 71. Ta-Chingoii region and Taman [Foggy] mountain. Habitat of sika, 

Manchurian wapiti, wild boar, and goral; an area in which tigers regulariy hunt. 
Here in the fog, the animal catches wapiti and sika coming out to sun themselves. 
Sudzukhin presei-ve, April, 1967. Photograph by V.E. Prisyazhnyuk. 

vegetation with an abundance of nut pine and oak. The animal also lives 
in pure nut pine forests, and also among mountains overgrown with 
deciduous shmbs, or in oak or nut-tree groves. Tigers only travel through 
dense spruce forests and do not remain in them. While tigers are attracted 
to river valleys and nut pine forests by wild boar, they mainly hunt abundant 
wapiti and moose in burned-over clearings (Kaplanov, 1948). Tigers are 
particularly attracted to rocky sections. "Permanent habitations are invariably 
found at places where mountain ranges are inaccessible, slopes very steep 
and precipitous, places with exteasive rock debris, overhanging cliffs, rocky 
crests, niches, and caves" (Baikov, 1925). Tigers can readily hide in rocky 
sections and, what is more important, there is less snow in rocky sections 
in winter. While hunting and wandering tigers prowl through quite high 
mountains, ascending to 1,600 m above sea level, nevertheless they prefer 
to move along gentle slopes and in low saddles. During times of movement 
the animal is seen in quite diverse biotopes, including sea coasts, hayfields, 


etc. Quite often they are confined close to villages, especially in winter 
when it is difficult to capture food. 

In the Primor'e and other parts of the Far East where frequent and heavy 
snowfalls occur, tigers avoid sections with a deep snow cover since the 
availability of ungulates in such areas is poor. Moreover, an unstable cmst 
hinders hunting. On thin crusts the tiger's movement is audible and frighteas 
wild animals; furthermore, the predator is apt to hurt its legs. The legs of 
a female tiger killed at the end of January, 1868 in the Ussuri region were 
131 severely injured. The fur ' 'on the inner side of the front and hind legs were 
worn away and the skin covered with bleeding sores" (Przheval'skii, 1870). 

The heavy bodied tiger moves with difficuhy in deep snow cover and 
breakable snow crust. The weight load per 1.0 cm^ of resting paw surface 
in Primor'e is equal to 158 g; among our wild cats this predator exerts the 
highest load on the resting paw surface (Formozov, 1946). In winters of 
abundant snow and snow crusts tigers suffer from hunger and sometimes 
even die of starvation. In the southern Ussuri region on March 1 (14) at 
a time of crusted snow blood was noted on the tracks of a tiger. This 
animal was evidently very hungry ' 'since it stopped along the way to eat 
a half-pecked rotten wolf-fish washed ashore in the previous autumn." Its 
feces "were small in quantity and contained . . . considerable undigested 
grass." Some days later the carcass of the tiger was found in a nut pine 

127 Fig. 72. Nut pine forest at source of a stream — habitat of tiger. Sikliote-Alin 

preserve. Photograph by V.K. Abramov. 


129 Fig. 73. Nut pine -spruce forest at the headwaters of the Ulakha — habitat of tiger. 

Here are also lynx, wild boar, wapiti and yellow-throated marten. Ussuri region. 
December, 1969. Photograph by A.G. Pankrat'ev. 

grove with no signs of wounds. The animal exhibited emaciation (Almazov, 

In the Far East, after winters of abundant snow accompanied by high 
mortality among ungulates, the food situation 8Ьаф1у worsens for tiger. The 
winter of 1914/1915 in the Sikhote-Alin range was extremely snowy. Depth 
of snow cover reached 100 to 150 cm. In that winter wild boar mortality 
was very high and over the next three or four years very few were sighted; 
in addition, hunters killed most of the weakened wapiti. A sharp reduction 
in number of ungulates the following winter resulted in the starvation of 
tiger and lynx and a decline in their population (Kaplanov, 1948). Near 
Temeya in 1915 an adult male tiger killed was not only emaciated but its 
intestines empty. It weighed only 70 kg versus a normal weight of 200 kg 
(Formozov, 1946). This predator had iaspected traps and devoured Siberian 
weasels caught in them, visited the camps of wintering hunters, and attempted 
to catch one of their dogs. In the snowy winter of 1935/1936, in the basin 
133 of the midcourse of the Iman, a young tiger weighing 110 kg, was found 
which had died of starvation. Its stomach was full of lichens (V. Abramov, 


130 Fig. 74. Broad-leaved forests with nut pine and fir — habitat of tiger. Suputin 

presei-ve, southern Ussuri region. September, 1970. Photograph by V.G. Heptner. 

In the Sikhote-Alin tigers suffered from starvation in 1957 and 1958. 
In these mountains wild boar, the main prey of tigers, had almost disappeared. 
The snow cover had been very thick for two consecutive winters, i.e., 
1956/1957 and 1957/1958, depriving wild boar of their meager food due 
to crop failure of pine nuts and acorns. An epizootic plague further decimated 
the swine. At the same time, the wapiti population was reduced to about 
one-eighth its former level (K.G. Abramov). Starving predators began to 
wander extensively; attacks on sika confined in parks, and even on horses 
and dogs, were reported. Ten deer were killed in a short period in a deer 
farm at Gamov in the winter of 1959/1960. In the same winter at Kras- 


130 Fig. 75. Coniferous forests along tlie middle Bikin — habitat of tiger. Ussuri region. 

Photograph by V.K. Abramov. 

norechka in the Chuguev region a female tiger broke into a courtyard and 
attacked a cow. In the Sikhote-Alin preserve tigers attacked horses on two 
occasions and several times stole dogs. 

Winter starvation has been a common phenomenon among tigers in 
Primor'e, since abundantly snowy winters are frequent there. Over the last 
110 years severe winters have occurred in 1855-56, 1876-77, 1877-78, 
1884-85, 1886-87, 1891-92, 1901-02, 1909-10, 1914-15, 1924-25, 
1925-26, 1926-27, 1932-33, 1935-36, 1938-39, 1940-41, 1941-42, 
1947-48, 1956-57, 1957-58, 1961-62, and 1964-65 (Nasimovich, 1950; 
Bromlei, 1963 and 1968). In the last 50 years, the winters of 1914-15, 
1941-42, 1947^8, and 1957-58 were especially bad for ungulates and tigers. 
Such winters have also been recorded for the Pri-Amur (1887-88, 1941-42, 
and 1942-43).''^ In the Primor'e region winters with abundant snow occur 

''^In the winter of 1970-71 in the Sikliote-Alin preserve wild boar were very scarce, 
and '"tiger tracks were far more numerous than boar". Evidently the predator smvived on 
Manchurian wapiti, which are more difficult for it to catch than wild boar. The tiger's situation 
was eased by the fact that the winter was not snowy; they were not stressed (E.N. Matyushkin). 
In the southern Ussuri region conditions were such that tigers struggled to live and in that 
winter some attacked cattle and even broke into cattle sheds. (V.H.) 


once in every two to nine years, and on the average once in every four 
years, sometimes consecutively for two to three years, and then tigers are 
especially hard hit. In the Primor'e exceptionally snowy winters recur once 
in six to fifteen years, on the average once in nine years. In such winters 
the snow cover reaches 70 cm in depth even by November instead of March, 
as is customary (Bromlei, 1963). 

Under unusual conditions in snowy winters, when an unbreakable ice 
crust is formed, hunting of ungulates by tigers is aided, since wild boar, 
wapiti, and roe deer slip while the predator moves well on such crusted 
ice. Tigers successfully preyed on ungulates in the Primor'e in the winter 
of 1877-78 because of ice crusts (Yankovskii, 1882). When moving through 
deep snow tigers quickly tire and often rest. Hence when the snow is deep 
or covered with a breakable crust, the predator sticks to rocky sites and 
sunny regions, and in moving from one area to another it often moves on 
the frozen bed of a river or stream, along wind- felled trees, pathways made 
by wapiti and other ungulates, or along man-made winter roads and ski 
tracks. If there is much snow on the ice, the animal tries to skirt it, choosing 
sections under coniferous trees where the snow is less. 

131 Fig. 76. Valley of Beloborodov spring with nut pine-broadleaved and nut pine 

forests along the slopes — site of tiger migrations along banks and valleys in the 
forest. Sikhote-Alin presei-ve. December, 1964. Photograph by E.N. Matyushkin. 


The Amur tiger is quite well adapted to low temperatures; it has a 
luxuriant, dense pelage and fattens by autumn. Cubs have been recovered 
which had spent the entire night in the forest at temperature of -40°C, but 
without apparent ill effects. From time to time, however, animals have been 
found with both eats, and even the tail, frostbitten. Evidently the main factor 
limiting the spread of tigers northward is deep snow cover (over 30 cm) and, 
consequently, the absence of wild boar, wapiti, and roe deer. In Traas-Baikal 
and the middle Pri-Amur, tigers were distributed up to 52° N. lat., i.e., farther 
north than at any other place in their range. The maximum depth of snow 
cover in this region does not exceed 20 cm and at some places is only 10 cm; 
its depth exceeds 30 cm only on the heights of the foothills (Rikhter, 1948). 
134 Food. In southeast Trans-Caucasus, the wild boar was the main prey 

of tiger. In the stomachs of tigers killed in the Lenkoran basin, only the remains 
of this ungulate were found (Satunin, 1914). Occasionally tigers preyed on 
roe deer, Caucasian red deer, and various domestic animals, including dogs; 
cattle were attacked only in winter (Dinnik, 1914; Vereshchagin, 1942). In 
Iran this predator preyed on the same species of animals, and also on goitered 
gazelle (Brandt, 1856). 

In Turkmeriia, Uzbekistan, and Kazakhstan wild boar was likewise the 
primary food of this predator. Bukhara deer occupied second place in many 
regions of Middle Asia, and roe deer second place in Kazakhstan. In 
Tadzhikistan, Bukhara deer have held first place in the food of tiger in recent 
years, with wild boar and domesticated animals coming second. For example, 
in Tigrovaya Balka in the lower reaches of Vakhsh River, from 1946 through 
1950 this predator attacked 19 deer, 17 cows, 4 horses, 5 asses, and 1 camel. 
In summer cows and horses straying from their herds or remaining alone 
at night became the prey of tigers in the tugais. The stomach of a tigress 
' caught on June 26, 1950 in a tugai at Kyzylsu contained the meat and bones 
of cows (Chemyshev, 1958). In floodplain reeds tigers lie in wait for goitered 
gazelle which visit water holes; the remains of gazelles killed by tigers were 
once often encountered in the reeds (Flerov, 1935). In Febmary, 1945 in 
Tigrovaya Balka the remains of goitered gazelle, wild boar, and deer were 
found in a den (Stroganov, 1959). 

In the lower Amu-Darya River tigers have sometimes killed and eaten 
jackals, chaus [jungle cats], and locusts (Pokrovskii, 1951). In Tadzhikistan, 
on the Vakhsh River, tiger feces often consisted of only locust remains, or 
an admixture of locusts and mole crickets, chitin of various beetles, hair of 
small mammals (mainly of complex-toothed rats [Nesokia indica]), and bird 
feathers (Rerov, 1935). In southem Tadzhikistan tiger feces sometimes 
contained the remains of mouse-like rodents, birds, and insects, and autumn 
and winter feces even contained kemels of oleaster {dzhida) and sea buckthorn 
fniits (Stroganov, 1961). 


132 Fig. 77. Broad-leaved forest. Habitat of tiger. Southern Ussuri territory. Photograph 

by V.K. Abramov. 

On the Zhana-Darya and around the Aral Sea in Kazakhstan, in addition 
to wild boar, tiger also caught saiga, goitered gazelle, wild horses, kulan, 
and mountain sheep (Ovis ammon arcat) (Meyendorff, 1826; Eversmann, 
1850). Goitered gazelles were preyed on during visits to water holes, and 
along the Hi (Alferaki, 1882). In reeds on the floodplains of the Syr-Darya, 
Chu, Hi, and other rivers, tigers evidently hunted saigas when they were 
numerous in these valleys during winter. In flooded areas and irrigation 
ditches tigers caught fish (sazan), which they eat with great enjoyment 
(Smimov, 1875). As in Tadzhikistan, in other regions of Middle Asia and 
Kazakhstan also, tigers attacked domestic animals, even camels, but most 
often dogs and horses. For example, in the winter of 1877 tigers killed all 
the dogs in a village beyond Chircliik at Kara Tyub (Smimov, 1879). "At 


places where cattle were abundant in Kirgizia tigers subsisted, especially 
in winter, almost exclusively on them" (Alferaki, 1882). In 1930 in southern 
Pri-Balkhash, in the region of Krasnyi town, tigers destroyed in a short 
period over 200 head of cattle, forcing the local populace in several places 
to migrate with their livestock to safer regions (N.Sh., 1930). 

Tigers formerly occupying Baikal and Trans-Baikal fed on wild boar, 
roe deer, wapiti, moose, and domestic animals, i.e., horses, cows, and 
reindeer. For example, the stomach of a tiger caught on November 22, 1953 
near Gazinur River contained the meat and hair of a roe deer; another killed 
on December 24, 1953 in the Mogochinsk taiga contained the remains of 
a wild boar. 

In the Far East, in Sikhote-Alin and other regions, tigers live on wild 
boar, wapiti, moose, sika deer, roe deer, musk deer, Manchurian hare, bear 
(brown or white-breasted*), lynx, wolf, badger, hazel grouse, and even fish. 
Their primary food has always been wild boar in these areas. From the 
stomach contents of tigers and the remains of their prey, 1 1 animals have 
been identified over the years; of these, 3 were Manchurian wapiti, 4 wild 
boar, 1 moose, 1 musk deer, 1 bear, and 1 lynx (Kaplanov, 1948). Tiger 
food in these areas (expressed as a percentage of total stomachs and carcasses 

132 Fig. 78. Nut pine-sprace forest at headwaters of the Ulakha. Habitat of a female 

tiger with her litter. Wild boar, wapiti, lynx and yellow-throated marten also are 
found here. Sikhote-Alin. December, 1969. Photograph by A.G. Pankrat'ev. 

*Refers to Eurasian black bear (Ursus thibetanus), which possessed a large white patch 
on the throat and chest — Sci. Ed. 


135 examined) comprised: wild boar — 40, wapiti — 30, moose — 10, musk 
deer — 10, bear — 6, roe deer — 3, and hazel grouse and other animals — 1 
(G.F. Bromlei). In Sikhote-Alin preserve wild boar were found 21 times 
(35.7%), wapiti 13 (22.1%), moose 6 (10.1%), brown bear 5 (8.4%), musk 
deer 8 (13.6%), roe deer 2 (3.4%), lynx 1 (1.7%), and hazel grouse 3 (5.1%) 
(V. Abramov, 1962). 

In years when the population of wild boar was low and their physical 
condition poor due to compulsory feeding on horsetails and twigs, the role 
of different species of animals as tiger food changed. For example, from 
1957 to 1959 the prey of tiger consisted of the following animals (based 
on number of carcasses found): wapiti — 20 (50%), wild boar — 12 (30%), 
brown bear — 2 (5.0%), sika deer — 2 (5.0%), and moose, roe deer, musk 
deer, and badger — 1 each (2.5%). Evidently in years of poor yield of nut 
pine and oak, and winters of abundant snow, resulting in emaciation of 
wild boar, tigers attack them less often and hunt mostly for Manchuriari 
wapiti and moose (K.G. Abramov; V. Abramov, 1962). From 1944 through 
1950, 17 instances of tigers attacking bears (brown and white-breasted) 
were recorded. 

Old and sick tigers usually confine themselves to smaller prey: piglets, 


Fig. 79. Remains of bull wapiti devoured by a tiger. Beloborodov spring, Sikhote- 
Alin presei-ve. February, 1968. Photograph by E.N. Matyushkin. 


young wapiti, roe deer, goral, and even hares and pheasants. For example, 
a lame tigress living in 1953 on the right bank of the Bir River, fed on 
hares which she caught on the island, and on wild boar piglets (Rodin, 1956). 

While prowling mountain streams and brooks, tigers skillfully catch 
fish by grabbing one in the water with a paw and throwing it on the bank. 
Turtles are scooped up when they surface for air. The infection of tigers 
by the helminth Paragonismiis westermanii, the intermediate hosts of which 
are fresh-water crabs and crayfish, proves that tigers eat them also (V. 
Abramov, 1962). Tigers sometimes eat pine nuts, shell and all, wild berries, 
fruits, and summer grass. Carrion is resorted to only in time of starvation 
(Baikov, 1925) (Fig. 82).* 

In the last century, when tigers were numerous in the Far East, their 
attacks on domestic animals, especially dogs, were comparatively frequent. 
Such attacks increased toward the end of winter. In the 1950's and 1960's 

136 cattle killed by tigers became a rare occurrence there. From 1956 through 
1959 in the Primor'e region, tigers attacked the following domestic animals: 
adult horses ten times, cohs twice, cows ten times, and calves once. In the 
same period eight sika deer were killed in a deer farm (V.K. Abramov, 
1962). Tiger attacks on domestic animals increase in years of a sharp 
reduction in population of wild ungulates, as occurred, for example, in 
1957-1959. Apparently domestic animals near or in villages, or left to pasture 
without proper supervision, fall victim either to young predators around 
three-years-old or to old and sick animals which cannot hunt for their normal 
prey (K.G. Abramov). 

137 In our countiy tigers pose almost no threat to man; man-eaters, famous 
in India, have never been known here, ahhough stray reports of attacks 
on man (see ''Practical Significance") have been noted. Man, however, 
has never been a 'source of food" for Soviet tigers. 

In adjacent countries tigers feed mostly on large animals. In the northern 
part of northeastern Cliina they hunt wild boar and wapiti, often attack dogs, 
and from time to time bear and small ungulates such as roe deer, goral, 
and catch hares and pheasants. The distribution of tigers in northeastern 
China (Manchuria) is generally associated with the availability of nut pines 
and wild boar: "Where there is nut pine, there is wild boar; where there 
is wild boar, tiger will also be found" (Baikov, 1925). 

In southern China wild boar, large deer, muntjak, domestic animals, 
and more rarely porcupines and pangolins serve as prey for tigers; tigers 
even eat frogs. In Yunnan in southwestern China their main prey are sambar 
deer, muntjak, and wild boar; on occasion they attack domestic livestock — 
pigs and buffalo calves (A.A. Sludskii). 

*Reference not clem-; may mean Fig. 81 — Sci. Ed. 


136 Fig. 80. Young wapiti in velvet killed by a tiger. Ta-Chingou, Sudzukhin preserve, 

May, 1966. Photograph by V.E. Prisyazhnyuk. 

In India tigers hunt for deer (sambar and axis), wild boar, buffaloes, 
nilgai, porcupines, peacocks, and domestic cattle; during floods they hunt for 
crocodiles, turtles, and fish. The stomach of some killed tigers were filled 
with locusts and mollusks. When starved, they catch frogs and mice (Jerdon, 
1874*; Bikhner, 1905; Pocock, 1939; Tate, 1947; Berten, 1954*). Tigers 
rarely attack mature male boar. They avoid carrion but sometimes when 
food is scarce they will consume rotting carcasses, including those of other 
tigers. Sometimes tigers lick saline soil (Pocock, 1939). (Methods of stalking 
used by tigers are discussed under "Daily Activity and Behavior".) 

It has been stated that a hungry animal consumes some 32 to 48 kg of 
meat in one sitting (Baikov, 1925). This is hardly possible, although it has 
been established that a sambar may be consumed by a tiger in two days and 
a buffalo in three days, leaving parts of the prey unconsumed (Corbett, 1957). 
In one sitting a tiger reportedly consumed almost one-third (30 to 40 kg) 
of a killed wapiti. The meat of the croup was consumed entirely and the 
bones of the legs and vertebral column gnawed roughly up to the middle. 

"Not in Literature Cited — Sci. Ed. 


The body of the wapiti was slit open across the chest cage. All the organs 
above this line (lungs, heart, and liver) remained untouched while the 
intestines and part of the stomach were consumed (V.E. Prisyazhnyuk). 
The normal daily ration of the tiger in zoological gardens is 10 to 12 kg 
meat (Obukhova and Shakhnazarov, 1949). 

In India, an adult tiger invariably kills no more than a single animal 
in one day. Young tigers, attacking a herd, may kill four or five cows in 
a single day (Jerdon, 1874*). In Sikhote-Alin an adult tiger kills and 
consumes per annum an average of 30 large animals of 100 kg (3,000 kg 
live weight). A family of tigers (mother and three cubs) consumed 280 kg 
meat in 20 days (wapiti 150 kg, wild boar 100 kg, piglets 20 kg, and musk 
deer 10 kg). In one year animals belonging to a single litter consumed about 
5,040 kg meat (Kaplanov, 1948). 

In connection with the fact that tigers sometimes sustain prolonged 
hunger, in favorable seasons a large accumulation of fat reserves is found 
in the abdomen, groins, and body cavities in layers 4.0 to 5.0 cm thick. 
Fat layers are also found between the muscles. A young three-year-old male 
caught on February 16, 1940 in Sikhote-Alin yielded 30 kg fat (Kaplanov, 
1948); a female caught on March 30, 1950 on the Pyandzha yielded 10.5 
kg fat and the layer in her abdominal cavity was 40 to 50 mm thick 
(Chemyshev, 1958). Thanks to fat reserves constituting 20% of overall body 
weight, tigers which have eaten nothing for even five to ten days remain 
content. Evidently they can withstand even longer periods of starvation. 

136 Fig. 81. Brown bear killed and eaten by a tiger. Tatibe River, southern Ussuri region. 

Autumn, 1961. Sketch by N.N. Kondakov from a photograph by V.E. Prisyazhnyuk. 

*Not in Literature Cited — Sci. Ed. 


Tigers drink water often and in quantity. In Middle Asia, especially 
in winter, they mostly employ rivers as water sources, and not lakes. They 
drink from lakes mostly in spring and summer, when the brackish water 
is less saline than in winter. 

Home range. A diet of large animals compels tigers to prowl widely in 
138 search of prey and often follow ungulate from one pasture to another. 
Therefore, the dimensions of the hunting territory of an individual are very 
large. In the Hi delta, where wild boar and roe deer are numerous, one tiger 
covered a distance of roughly 50 cm x 20 cm along a stream and hunted 
periodically in different sections of this area. As soon as the number of 
pigs had been shaiply depleted, the predator deserted the region and moved 
into another several hundred kilometers away from the former. In Tigrovaya 
Balka on the lower Vakhsh in Tadzhikistan, which ab()unds in ungulates, 
the territories of three tigers, though overlapping at places, covered an area 
roughly 6.0 to 7.0 km in diameter per animal (Flerov, 1935). In this same 
region, according to other data, the individual territory of a tiger usually 
does not exceed 30 to 40 km^. However, tigers do not live for long at one 
place and in a few days or weeks move into an adjacent region or one farther 
away (Stroganov, 1961). 

In Sikhote-Alin preserve in the winter of 1939 to 1940 a single tigress 
periodically visited a whole system of rivers and ranges and was encountered 
in a territory 60 km x 70 km. An adult male, also in winter, was periodically 
sighted in a section 80 km x 40 km (Kaplanov, 1948). In this same preserve, 
in places rich in ungulates, tigers were sometimes confined to a territory 
around one or two springs covering an area of 400 to 500 km^ (G.F. 
Bromlei). In wintei-s of heavy snow the territory regularly visited by tigers 
asually decreases, and the animal beats pathways along which it prowls. 
It may be assumed that tigers in summer spend a more settled way of life; 
yet even then they often move 10 to 30 km. A tiger prowls throughout 
its territory in a rather circular manner and hence is periodically sighted 
at the same place at intervals of a week or month, depending on the size 
of the animal's domain. 

As in the case of other species of cats as well as bears, tigers demarcate 
their territory with characteristic markings; the animal rises on its hind legs 
and scratches the bark of trees with the claws of the forepaws (Baikov, 
1915; Allen, 1938; Arsen'ev, 1950). The bark on a tree is sometimes stripped 
off to a height of 250 cm. From such markings the dimensions of the territory 
of a single animal can be judged. Special scent marks in the form of urine 
and excreta (tigers have preanal scent glands) are also deposited. For this 
purpose particular stones or trees are selected and periodically sprayed with 
urine. On finding a suitable site, the tiger sniffs it, then deposits its urine. 

Before undertaking a hunt a tiger will usually move along the top of 


the mountain chain, from where it can inspect both slopes; often it rests 
on a high rocky outcrop overlooking a river valley and views the surround- 
ings. '"Having killed and eaten its prey, either totally or partially, the animal 
roams its selected territory. At first it rolls in the snow every 100 to 200 
m, leaving spots of blood in the print of its body, which reveal its departure 
from a recent kill. During the day the animal often rests on its abdomen, 
urinates, defecates, and paws the snow and ground quite often, moving tens 
or even more kilometers before lying down in a clear sunny section devoid 
of trees. At night, on the contrary, it selects for rest a dense spmce thicket 
which easures a few degrees more warmth than a sparse growth' ' (Kaplanov, 
1948). After resting, which may continue for 12 or more hours, the animal 
resumes its journey, only rarely stopping to roll in the snow. During long- 
distance forays, periodically stopping and resting, sometimes however 
without stopping, a tiger may travel for several days and cover some tens, 
even hundreds of kilometers. In a single day the animal may traverse 20-50 
(Kaplanov, 1948) to 100 km (Baikov, 1925). Only during estnis do the 
female and male stay together in a comparatively small area. Tigers prefer 
an area rich in wild boar or Manchurian wapiti. 

Unlike single animals, a tigress with cubs less than two years old will 
occupy a very small territory. For example, in Sikhote-Alin, in a section 
rich in ungulates, a tigress with cubs lived in an area 5.0 km x 3.0 km from 
December 24, 1940 through January 15, 1941 (Kaplanov, 1948). A tigress 
with cubs aged 2.0 to 3.0 years will extend her territory much beyond that. 
139 In 1957 in the upper reaches of the Maikhe River a family consisting of 
a tigress and two cubs in their second year was discovered. Until the spring 
of 1958 this family lived in a fairly limited territory. Later they enlarged 
the zone of their activity and by autumn of 1959 occupied an area of 50 
km X 60 km (V. Abramov, 1962). 

Burrows and shelters. Solitary tigers usually do not establish a perma- 
nent den but lie down to rest close to their kill. In the river valleys of Middle 
Asia and Kazakhstan favored sites for rest during the day are the edges of 
extensive reed beds adjoining a tugai or open forest glade. The den is usually 
situated in the shade of a single oleaster {dzhida) or turanga tree [Euphrates 
poplar] occurring among reeds or reed grass [veinika], which becomes 
strongly trampled down. Dry grass serves as a bed. At times a very distinct 
path to the den is apparent in trampled grass, adjacent to an area 30 m x 
40 m^ of trampled or beaten grass scattered with bones, bits of skins, and 
wool discarded by the animals. Obviously the stench from such areas is 
hardly pleasant. Judging from trampled grass the animals undoubtedly rest 
in several places, moving with the shade (Stroganov, 1961). 

In the Far East tigers select for rest a fairly concealed place somewhere 
under a nut pine, sometimes even on sunny sands or rock. They return fairly 


regularly to favored sites and make a den. In winter they rest directly on 
the snow. A tigress hides her cubs in rocky areas of the mountains or in 
dense vegetation, selecting a niche in the rocks, cracks in a precipice, or 
a cave for a den. 

In Middle Asia and Kazakhstan dens for cubs are located in dense 
galleries of reed or in tugai, on an islet unaffected by water. The den is 
established in a natural soil depression and sometimes covered with dry 
leaves and grass, but usually lacks bedding. Often the den is well concealed 
by dense reeds, wild vines, lianas, and other plants. A tigress with very 
young cubs will approach her den cautiously and attempt to leave behind 
as few traces as possible. 

Dens are established in areas of abundant wild boar and other ungulates. 
A tigress uses the same den for several consecutive years. The den of a 
dead tigress is often taken over by another female. When the cubs grow 
up and begin to follow their mother, she leaves them behind in a temporary 
den nearby, within 500 m, when going out to hunt. Hunting is done stealthily 
so as not to frighten ungulates wandering close to the den (Kaplanov, 1948). 
In India tigers set up dens under rocks or in caves. On one occasion, however, 
a pair of cubs was found beneath a prickly bush (Jerdon, 1874; Pocock, 
1939). Around the den scattered bones and excrements often exude a 
characteristic strong odor. However, within and around the den per se, as 
long as her cubs are small, the mother le<ives neither bones nor other remains 
of prey (Anderson, 1964).* 

Daily activity and beliavior. Within the Soviet Union tigers become active 
around twilight; they may hunt at any time of the day but most often after 
sunset, in the first half of the night, or later at dawn. Tigers cannot withstand 
heat well. In India they usually set out to hunt at sunset and hunt throughout 
the night, slowly covering their territory along trails (Pocock, 1939). 

Tigers employ several methods for hunting wild boar, wapiti, moose, and 
other animals, but mainly resort to ambush. On finding the track of a wild 
boar or wapiti, the predator follows the spoor until it comes fairly close to its 
prey. Here it turns to the leeward side and concealing itself on the path awaits 
the approach of its intended victim. The olfactory sense is rather poor in 
tiger but nevertheless the animal is capable of tracking a quarry even when 
the spoor is an hour old. The predator locates its quarry mainly through 
the senses of audition and vision. Hunting tigers are assisted by the bright 
and variegated color of their coat, which blends with the surroundings. When 
a tiger runs through the forest among shrubs the black, yellow, and white 
colors of its coat merge and the animal appears a monochromatic brownish- 
gray (Arsen'ev, 1949). When hunting in tall reed thickets tigers sometimes 

*Not in Literature Cited — Sci. Ed. 


rear up on their hind legs or leap upward in order to inspect the surroundings 
(Khakhlov, 1928). 
140 In summer tigers lie in wait for wapiti, moose, and roe deer at back- 
waters, where these ungulates go for food, and at ponds, and also at salt 
licks; in winters of heavy snow tigers maintain a vigil along their trails. 
The predator approaches the quarry closely in one or two bounds and throws 
it to the ground by striking it with its forelegs. The toppled prey dies almost 
immediately due to cervical vertebrae broken at the occiput or a broken 
occiput per se. A wapiti killed in Sidzukhin preserve had a broken neck; 
on the right shoulder, struck by the tiger's forepaws, the skin was torn and 
showed the imprint of all five paw cushions, the same as a pug mark in 
mud (V.E. Prisyazhnyuk). Huge canines help the tiger kill an animal quickly. 
It requires more time only for large wild boars, bears, and bulls. When 
dealing with big game tigers attack from the sides and rear. Having felled 
its prey the predator does not catch hold of it immediately but makes a 
few more strikes or pounces. A tiger will give up the chase after pursuing 
a potential quarry for about 100 to 200 m. 

Tigers have adopted a unique method of hunting in the Ta-Chingou 
area in the Sudzukhin preserve (Figs. 71, 80 and 84). There wapiti and sika 
deer regularly wander onto the flat sandy shores of the bay at night and 
sometimes even in the day. On the night of May 23* the fog was heavy. 
Taking advantage of this fact a tiger approached within 10 to 12 m of a 
herd of seven wapitis standing right at the edge of the water. Launching 
its attack from behind a small undulation, the predator overtook one of the 
velvet antlered wapitis [Fig. 80] in three short bounds. Later, it dragged 
the carcass half-a-kilometer into a grove of hazel and young oak trees. 
Hunting by the sea in fog is the usual method employed by tigers in this 
region. The fog is so thick that a human is not visible until approached 
to 20 to 30 m. Tiger tracks in such mists on coastal sands right at the water's 
edge sometimes cover a kilometer. Spoor are visible there both in summer 
and winter (V.E. Prisyazhnyuk). 

Having tracked an animal, a tiger will not run long after its quarry. 
The only exception is the wild boar, which a tiger chases with tenacity; 
the predator will systematically kill one after another until the entire herd 
is destroyed. Subsequently, it will set off to find a new herd. Hunters refer 
to this method of tiger hunting as "grazing" the pigs. 

In attacking animals a tiger makes very little noise and hence does 
not scatter a herd. In Sikhote-Alin preserve a single tigress stole one animal 
after another from a herd of wild boar, destroying the lot in just a few days. 
In spite of repeated attacks boars still remained in the same section of the 
preserve. Shortly after the tigress, a male sneaked up on another family of 

*Year omitted in Russian original — General Editor. 


wild boars, made a couple of pounces, and catching one of the pigs carried 
it in bounds 200 m away into a bush. Having consumed the catch, it repeated 
this tactic a couple of days later. The pigs in the herd, though uneasy at 
night, appeared calm and grazed quietly during the day on the sunny slopes 
of the river. On another occasion an attack by a tigress on a resting female 
wapiti was observed. The attack was so swift that the deer had no time 
to even stir from its bed. 

In December, 1941 -January, 1942, working at the site of permanent 
tiger occupation in the basin of the Fata and Sitsa rivers (Terneya region), 
G.F. Bromlei (1964) did not sight even once the exodus of wild boar due 
to their decimation by this predator. Yet he found the remains of slaughtered 
pigs in several places. From the traces it was established that the tiger hunted 
at night. Sneaking to within 10 to 12 m of a herd of pigs, it executed two 
or three bounds, caught hold of the victim, and carried it away without 
shedding so much as one drop of blood. Only from a narrow trail on the 
snow, caused by the dragging legs of the prey, was Bromlei able to establish 
that the predator had succeeded in its hunt. The remaining boars, frightened 
by the attack, scattered from 200 to 300 m and then settled into a fresh 
shelter for the night. 

In the Soviet Union this predator is relatively silent during a hunt. 
Finding himself during the course of one month in a region where tigers 
live, G.F. Bromlei never heard a tiger voice even once. In India, contrarily, 
during a hunt and after it, a tiger roars frequently (Corbett, 1957). 

A tiger will even tackle a bear, sometimes one much larger than itself. 
Having tracked its victim, the predator selects a rock or wind felled tree from 
141 which it can ambush the quarry from the leeward side. It springs upon the 
approaching bear from an overhead position, catches it by the chin with 
the claws of one forepaw and the throat with the claws of the other, and 
clamps its jaws on the spine. Bears are generally afraid of tigers and, coming 
across their tracks, run away perpendicular to the trail. Confronted with 
a tiger, a bear will try to escape attack by climbing a tree (Baikov, 1925). 
On February 6 in Sikhote-Alin a tigress discovered the den of a female 
bear with yearlings. Digging up the den from the side opposite to the 
entrance, she proceeded to harass the mother from both sides alternately. 
Eventually the tigress seized the bear from the rear with her claws, behind 
the legs, dragged it out of the den, and quickly killed her by clamping its 
jaws around the cervical vertebrae at the occiput. The cubs, weighing 30 kg, 
each, she killed within the den (Kaplanov, 1948). In the Ussuri territory 
the hair of bears has been found repeatedly in the spring and autumn feces 
of tigers (April 24, May 7, and November 10, 15, and 27). Eariy in May, 
1951 on the bank of Tatibe River (Iman tributary) a bear was found (body 
length 158 cm, weight 170 kg), which had obviously been mauled by a tigress 


(Fig. 81). All the fatty parts of the body had been devoured — the back, 
hams, and fatty layers in the groin region. Fecal deposits, urine trails, and 
three separate lairs within 10 m of the carcass of the bear in which the 
tigress had laid up for three or four days were found. 

In the upper reaches of Pfusung River (west of Ol'ga Bay, Primor'e 
region) in September, a small white-breasted [Eurasian black] bear climbed 
up and down a tree several times trying to save itself from a tiger. Evidently, 
the tiger harassed the bear a long time since the bark of the tree was totally 
peeled off at places. In 1949 at the end of December, in the basin of the 
lower Sinanche (Iman tributary), hunters chasing a family of tigers found 
a white-breasted bear hiding from one in a tree. Judging from its badly 

148 Fig. 82. Track of a tiger (male) traversing deep snow witii impression of its tail 

also visible. Beloborodov spring in Sikhote-Alin preserve (Sitsa basin). February, 
1968. Photograph by D.N. Matyushkin. 


mauled skin and tufts of detached fur and other traces on the snow, the 
bear had been wounded by the tiger's claws and remained in the tree for 
over a day. 

Brown bears fall victim to tigers more often, since white-breasted bears 
usually live in hollows or among close-set rocks and are thus less accessible. 
In the middle Sikhote-Alin only three instances of killing of white-breasted 
bear by tigers have been recorded over several years of observation. 

Over 15 instances of tiger attacks on brown bears resting in dens located 
on the slopes of the middle Sikhote-Alin (Sikhote-Alin preserve) were 
recorded from 1952 to 1959, mainly in late autumn and early spring. Tigers 
did not always succeed in killing their quarry. The largest of the bears 
escaped from the tiger's claws, and having been chased from their dens, 
began to wander in the taiga, becoming "rovers". From 1940 the number 
of tiger attacks on bears has risen steadily, due, no doubt, to a general 
reduction in ungulate population. Evidently in the absence of ungulates tigers 
resort to feeding on bears (Bromlei, 1965). 

As mentioned above, a tiger is not always the victor in an encounter 
with a bear. In the winter of 1959/1960 a tigress lived with her two-year- 
old cubs in a ravine in the Sikhote-Alin. Once, having killed a wild boar, 
she left it with her cubs and went away. A large bear wandered into the 
area and on her return to the den the tigress attacked it. During the ensuing 
contest the bear killed the tigress. In spite of serious wounds, the bear 
proceeded to eat the wild boar and then the tigress (Sysoev, 1960). In 1913 
on the Bol'shoi Sinanche River a large brown bear knocked down a tiger. 
In 1960 in Sikhote-Alin preserve a young tiger became the prey of a bear 
(V. Abramov, 1962). A similar incident has been reported in the Pri-Amur 
(Krivopusk, 1957; Fig. 88). 

A hungry male or a lone tigress, after killing a large animal weighing 
100 to 150 kg, eats heavily and rests in the vicinity for five to ten days. 
Usually a tiger remains for only three or four days by its prey, until the 
meat begins to freeze solid or to decompose. 

Sometimes, having killed an animal, the tiger leaves it untouched and 
returns to it later. In such an event it sometimes covers the kill with brush, 
snow, and grass which it uproots with its teeth. It consumes large prey skin 
142 and all, leaving only the feet, hooves^ and head. It even cracks the long 
bones of moose. Having killed an animal the tiger usually drags or carries 
it into a bush close to water. After feeding it rests for much of the day, 
lying on its side with the legs outstretched, getting up occasionally to drink 
from the water hole. If hunting has been unsuccessful, it may continue to 
hunt even during the day. 

Like other species of cats, tigers are very clean animals. They often 
lick their coat and paws. After a hearty meal they love to roll on the snow 


and in summer bathe in a river, submerging the muzzle in water and 
"gargling' ' to rinse the mouth free of remnants of meat and blood. Claws 
are cleaned by stripping soft bark. On hot days, as a defense against blood- 
sucking flies, they lie in water with only their head protruding, or stretch 
out in the spray of a waterfall. 

Tiger movements are graceful, quick, and nimble. They can creep 
through grass only 60-70 cm high and remain completely undetected. The 
animal's pace is usually 50-80, and up to 100 cm long; it begins to sprint 
only when chasing its prey and may then leap as far as 4.0 m. The maximum 
length of a leap is 6.0 to 7.0 m, and a height of 2.0 to 3.0 m. Tigers negotiate 
precipices and rocks very well but generally cannot climb a vertical tree 
unless it has close-set branches. Exceptions are known, however. A tigress 
pursued by hunters clawed up a tree devoid of branches to a height of 7.5 
m (Pocock, 1939). Tigers are excellent swimmers, and large rivers (Amu- 
Darya and Amur) or sea channels up to 5.0 miles wide pose no problem. 

Adult males and single females not in estrus live alone. A tigress with 
growing cubs aged 1.0 to 3.0 years moves with her offspring; the latter 
sometimes comprise cubs from two litters. This explains why tiger families 
of four or five animals are sometimes seen. When tigers were abundant 
in the Far East, groups of even 7 to 13 animals were recorded (Kaplanov, 
1948). If such groups actually existed at all they probably represented two 
families which for some reason had joined together. 

The tiger possesses tremendous strength and is capable of carrying off 
animals far larger and heavier than itself such as horses, cows, and buffaloes 
up to three years of age (Brandt, 1856). A tiger can transport a horse or 
cow teas and even hundreds of meters from the place where it killed, it being 
carried, not dragged. Corbett (1957) tracked a tiger which carried a fully 
grown cow 4.0 miles (7.5 km) away from the place of its kill. Another tiger 
dragged a killed bull 13.5 meters. The bull could not be moved by 13 men. 
A tigress was seen swimming across a river holding a cow in her mouth 
(Pocock, 1939). Once a tiger jumped into a pen, killed a calf, and catching 
it in its teeth leaped back to a rock situated 3.6 m above the pen's level 
(Allen, 1938). If the prey is so heavy that the tiger cannot carry it, it is 
often abandoned. When the predator carries its prey by the neck, the rear 
of the body drags, leaving behind a distinct trail. If, however (as for example, 
a deer), the tiger holds the kill by the middle of the back, the portage trail 
may be faint or nonexistent. A tiger can carry a large wild boar in its teeth 
through dense mshes, jump onto rocks, and even climb a steep slope. 

This predator is less afraid of humans than other large animals, but 
nevertheless usually moves away from them. Tigers usually initiate an assault 
only when injured or chased, confronted by dogs, being approached at a 
den with cubs, or in an unexpected or chance encounter at night. Tigers are 


generally more aggressive at night and approach man more boldly (Kaplanov, 
1948). A tigress will sometimes attack to protect her cubs. Ussurian hunters 
who had captured several dozen cubs reported only one instance when a 
tigress, frightened off by shots fired in the air, returned to her litter and 
had to be killed. A tiger does not attack when men approach its kill (also see 
section "Territory" for further information on the behavior of tigers). 

Tigers caught young respond well to training. Even adult animals can 
be trained. 
143 Seasofial migrations and tfansgressions. It was noted above that single 

tigers wander much of the time in search of prey in a definite and fairly 
large region. Sometimes these wanderings are regular. For example, in 
Kazakhstan and the Far East tigers in spring climb high into the mountains, 
following grazing ungulates, and descend to lower altitudes in autumn. 

Periodic emigrations of tigers are definitely associated with a reduction 
in population of wild boar, or their migrations, or fires in the taiga, and 
in large sections of floodplains, and tugais. 

In Tadzhikistan it has been noted that an increase in the tiger population 
is directly related to massive fires in the tugais on the Afghan banks of 
the Pyandzh. In the lower reaches of the Syr-Darya tigers have been sighted 
only twice in the last 30 years, once in 1926 and again in 1945, and on 
both occasions were trailing herds of wild boar moving along the coast 
of the Aral Sea from the delta of the Amu-Darya. On the latter occasion 
they covered over 1,000 km calculated in a straight line. Reportedly, tigers 
following wandering wild boar in winter passed through the Kyzylkum. 
In 1930 in the Hi delta, consequent to large-scale slaughter of wild pig for 
meat, the tiger population dropped sharply. As a resuU, tigers emigrated 
from this area that year. Two moved upstream along the river : one was 
killed 500 km from the delta and the other entered China along the valley. 

Tigers caught in the last century in central Kazakhstan and also at Biisk 
and Barnaul were intruders who had left their permanent dwelling areas 
(Balkhash and Ziiisan) located at a distance of 400 to 600 km in a straight 
line.^^ In southeastem Trans-Baikal in the last century tigers were sighted 
only periodically, once every five to eight years. At that time they were killed 
near the Argun, Gazimur, and Shilka rivers. Following reindeer, the predators 
intmded into Stanovoi range from the upper reaches of Amur and Shilka 
(Brandt, 1856; Sel'skii, 1856). In southeastern Trans-Baikal they were seen 
from time to time early in the present century as well as in the 1950's and 
1960's. In 1953 about eight tigers were reportedly seen there (A.N. Leont'ev). 
One was killed on November 22, 1953 at Gazimur River in the Ust'-Kara 

^^Recognizing this habit of tigere — to wander or undertake extensive migrations, 
Kazakhs popularly call them '"road" or "traveling" leopards. 


region of Chita district, and another in the same year on December 24 on 
the bank of Shilka in the Mogochinsk region (Sludskii, 1966). From 
1963-1966 tiger tracks were detected repeatedly along the Uryumkansk River 
in the Plyiisnino region and along the Gazimur River near Kiingir, 
Kurumdyukon, Kaktolg, and Kachugai stations (Sinitsyn, 1966). Evidently 
they entered Trans-Baikal from the Great Khingan mountain range in China 
or from the Amur region. 

A tiger caught in 1905 close to Ust'-Maya (Aldan; Ognev, 1935; 
Galchenko, 1959) had traveled 1,000 km in a straight line from its region 
of permanent residence. Animals seen from time to time in the Verkhoyansk 
range (Silant'ev, 1919) and crossing the Stanovoi range up to 56° N. lat. 
had traveled no less distance. As mentioned above, they used to move that 
far north in pursuit of migrating wild reindeer. 

In the Far East, around 1886, tiger migrations were recorded for two 
consecutive winters moving west to east, from the Sungari in China to 
Shikote-Alin. All of the tiger tracks ran eastward according to Arsen'ev 
(1949). A similar migration was observed in 1913 (Kaplanov, 1948). There 
are records of annual migrations of tigers from northeastern China 
(Manchuria) into the Soviet Union, the region of Bikin River and other 

In 1924 hunters came across the track of a large tiger and tracked it. 
The chase continued for 22 days and in that period the tiger had moved from 
the Tetyukha to Taukha rivers, covering a distance of 1,000 km. Later the 
tiger crossed into the Suchan valley and from there to the Daubikhe River, 
where its trail was again picked up by the hunters, who followed it for another 
15 days up to the Iman (V. Abramov, 1962). In earlier years tigers intraded 
almost every autumn for a few days into the Suputin preserve on the Suputin 
144 River (southern Ussuri region), descending from the Dodyan'-Shan mountain 
to the sources of Kamenka and Anikinsk springs. At other times of the 
year their tracks were never seen in the Suputin preserve (Bromlei and 
Gutinkova, 1955). 

In the last century tigers were reportedly seen on Sakhalin [Island] in the 
Tym'-Poronai valley in the upper reaches of the Тут' and in the mountains 
near the Poronai River (Schrenk, 1858; Shmidt, 1868; Przheval'skii, 1870; 
Chekhov, 1956). Information is available, which requires confirmation, that 
tracks of a tiger were encountered on that island in 1950 and 1965. Tigers 
could have reached Sakhalin by swimming the 7.0 km width of NeveFsk 
Strait. Because of the very deep (3.0-4.0 m) and porous snow cover, a 
permanent residence for tigers on Sakhalin Island is impossible (for other 
tiger transgressions see "Geographic Distribution"). 

Tiger migrations into the Soviet Union from the Korean Peninsula have 
also been observed. At the end of the 1890's tigers crossed on ice over the 


Amlokan' River into the Korean Peninsula from northeastern China 
(Manchuria) (Von Hon Gu). 

Reproduction. Estrus, gestation, and parturition among tigers of northern 
subspecies, according to the latest observations in the Far East and in zoolo- 
gical gardens, are not restricted to any particular season. Newborn cubs 
may be found at any time of year. For example, in December, 1932, a 
month-old cub weighing about 2.5 kg was caught in the Ussuri region on 
the Baitsuka River, the right tributary of the Iman. In January, 1933, four 
dead cubs weighing 4.0 to 6.5 kg were found on the Iman River. In May, 
1933 a female tiger was caught near the Sanchekheza River with five fully 
developed fetuses in her womb. In the winter of 1937/1938 three cubs aged 
three or four months were found on the upper reaches of the Iman (Salmin, 
1940). Small cubs were found in this same region on January 9, April 13, 
July 22, October 19, and December 7 and 13 (G.F. Bromlei). In March, 
1957, a litter of four cubs aged 1.0 to 1.5 months was found (K.G. Abramov). 

A female caught on March 30, 1950, in southwestern Tadzhikistan at 
Pyandzh was in heat and had recently mated with a male as detected from 
spermatozoa present in the swab of the uterine cervix. A second female, 
caught along the Kyzylsu on June 26, 1950, was lactating (Chemyshev, 
1958). Cubs have been sighted in Kazakhstan from June to August (N.A. 
Severtsov, 1861). Early in February, 1909 feces of four cubs roughly one- 
month old were found on the lower reaches of the Hi. In early April, 1908, 
on the shore of [Lake] Balkhash in the region of the lower Ayaguz three 
small cubs were caught (Shnitnikov, 1936). 

In Soviet zoological gardens tigers in heat can be seen at any time 
of year, once every two or three months, but sometimes every month, or 
only twice a year. Three Ussuri tigresses in the Moscow Zoological Garden 
gave birth as follows: one ("Chizhik") twice in March, once in April, twice 
in May, and once each in October and November; another ("Naya") once 
each in May and June; and the third ("Lyuba") twice in May (Afonskaya 
and Krumina, 1956). 

Births in zoological gardens occur most often in two seasons — spring 
and late autumn; however cubs usually appear in May (46% of all newborns). 
Thus the periods of heat formerly reported for different regions of the USSR 
(January and FebRiary in Trans-Baikal, December and Januar}^ in Kazakhstan, 
and end of December and January in the Far East) are only partly correct. 

In the northeastern part of China newborn cubs are usually seen in 
April to June but most often in May. In southern China small cubs are seen 
not only in spring and summer but also in winter. In southwestern China 
(Symao, Yunnan) cubs are more often seen in April to June and September 
to October. That Indian tigers have no definite breeding season has long 
been known (Jerdon, 1874). The absence of strict seasonal periodicity in 


the appearance of litters among tigers points to their southern origin. 
Nonseasonal reproduction is characteristic of many tropical animals. 

Estrus among tigresses continues for 12 to 18 or, rarely, 25 days. Among 
Ussuri tigresses in the Moscow Zoological Garden, it extends for 3 to 13 
145 days (Afonskaya and Krumina, 1956). A female in heat makes a charac- 
teristic nimble, sniffs frequently, micturates often, rolls on her back, splays 
her legs, and plays with the male. She eats little or no food. During this 
period she usually moves with a single male, but in areas where tigers are 
numerous, two to six males follow a single female. During heat tigers roar 
more often than at any other time. The female attracts males by her roar. 
Serious fights occur among males. They also become less wary during rut. 

The mating period of tigers extends for 5 to 8 days, and sometimes 
up to 18. According to observations made in the Moscow Zoological Garden, 
up to 20 matings occur in a day, with an average of 11, during the mating 
season (at night the animals separate). The interval between successive 
matings may vary from three to four minutes to several hours. In all, 43 
to 123 matings have occurred in this period. While mating, the tigress lies 
on her chest and elevates her rear; the male grips the nape of her neck in 
his teeth (Afonskaya and Krumina, 1956). 

According to observations in zoological gardens, the duration of 
gestation is 98 to 112 days (Satunin, 1915; Baikov, 1925; Pocock, 1939; 
Shereshevskii, 1940). Commonly, gestation requires 105 days. Among 
Ussurian tigers gestation extends for 95 to 107 days, with an average of 
103 (calculated from the first to the last mating). At the end of estrus the 
female isolates herself from the male. In zoological gardens after the mating 
season males and females have to be separated immediately since fights 
ensue between them. The male does not take part in raising the young. 

Tiger litters throughout the USSR consist of two to four cubs, rarely one, 
and even more rarely five or six. Cubs die quite often in the first few months 
due to various causes and hence two or three are usually seen with the mother 
and more rarely one or four. In different parts of the range in the Soviet 
Union the following number of cubs have been recorded in a litter; Trans- 
Caucasus — two litters with two cubs each; Kazakhstan — three with two 
each, one with three, and one with four; Tadzhikistan — two with one each; 
and in the Far East — seven with one each, seven with two each; eight with 
three each, and two with four each. Additionally, a female with five fetuses 
was caught in the Far East. I.T. Trofimova, the tiger trapper, came across 
a female with four two-year-old cubs in 1953 in Sikhote-Alin near the 
Tudo-Vak River. From 1936 through 1957 hunters found five litters with 
two cubs each, ten with three each, and two with four each. From 1957 
to 1959, four litters were registered with one cub each, eight with two each, 
and three with three each (V.K. Abramov, 1962). Among Ussuri tigers in 


the Moscow Zoological Garden the number of cubs in a litter is the same 
as in the wild. Most often two cubs occur in each litter (50% of all cases) 
but the fertility of various females is not identical. The tigress "Chizhik" 
usually gave birth to three cubs (Afonskaya and Krumina, 1956). In the 
Peking Zoological Garden in 1958 there were either two or three cubs in 
the litters of two Ussurian tigresses. In thesouthem part of the range, for 
example, in the South China tiger in Yunnan province usually one or two 
cubs occur, rarely three or more, although a case is known here of a female 
captured with six embryos (A.A. Sludskii). Other southern China tigers 
produced two cubs (twice) and four cubs (twice) (Allen, 1938). In India 
two cubs are common, more rarely three or four, and very rarely five or 
six (Jerdon, 1874; Pocock, 1939). Tigresses with seven embryos have also 
been reported (Beriff, 1932).* 

Growth, development, and molt. Cubs are usually bom blind and 
helpless but sometimes with their eyes open (evidently when gestation is 
greatly prolonged). The weight of newborn Ussuri cubs varies from 785 to 
1,043 g, body length 31.5 to 40 cm, tail length 13 to 16 cm, and length of 
ears 1.5 to 2.5 cm. The claws are not pigmented. Cubs born blind gain sight 
on the sixth to the eighth day (according to other data on the fifth to the 
tenth day). The ears open on the fourth to fifth day and the young begin to 
respond to sounds at some distance by the twelfth to the fifteenth day. There 
are no teeth at birth but the places where they will appear are distinct on 
the gums. Cubs cut the two medial incisors on the 13th day. Two incisors 
are seen on both the upper and lower jaws on the 23rd day, six incisors 
on both the upper and lower jaws on the 42nd day, and upper and lower 
canines on the 53rd day. There are two cheek teeth, two canines, and six 
146 incisors on the upper jaw on the 63rd day. At the age of 8.5 months the 
cub sheds the middle pair of incisors and the other incisors at the age of 
9.0 months. Molt of the juvenile fur coat occurs at the age of 3.5 to 5.5 
months (Afonskaya and Krumina, 1956). The color of the juvenile coat is 
the same as that of adults but the background much lighter and the stripes 
light brown. The ears of small cubs are relatively large. As in adults, a 
bright black color with a white spot in the middle occurs on the back of 
each ear. Therefore, this signal color is fully developed at an eariy age. 

Cubs grow and develop rapidly. On the 12th to the 15th day they begin 
to crawl around the den and by the 20th to the 30th day emerge, walk on 
the snow, and climb trees. Even two-year-old cubs weighing up to 60 kg 
may climb trees, but most lose this ability early. 

The growth rate of young tigers can be judged from the weight chart 
of "Sirotka", a cub fed artificially in the Moscow Zoological Garden: 

*Not in Literature Cited — Sci. Ed. 


























When 35 to 36 days old cubs begin to lick meat and in 43 days attempt 
to eat it but receive no encouragement from their mother. After 48 days 
cubs tear off bits of meat. At the age of two months they begin to eat meat 
regularly but the mother continues to suckle them to the age of five or six 
months. The mother transports two-month-old cubs from one den to another. 
Leaving them there she sets off to hunt, and having caught a prey, takes 
her offspring to it. The litter remains near the kill while she again sets out 
to hunt. As long as the cubs are small the mother leads them to each kill. 
She does not leave her offspring of less than six months of age alone for 
long; those older than that she leaves alone for five or six days and two- 
year-olds for up to fourteen. Very young cubs left alone in winter sometimes 
freeze to death (Salmin, 1940; G.F. Bromlei). When prey is plentiful the 
litter is not moved far and remains in a relatively small area. 

Cubs are extremely agile and playful. When their hunger has been 
appeased they romp continuously and play with their mother until they feel 
sleepy. In play one cub will hobble side- ways, hide pretentiously, and wait 
for another cub to discover it; it then jumps out and tries to pin the other 
down by catching hold of its hind legs or tail. Cubs play with the mother 
in the same way. Sometimes a cub having caught another by its tail with 
its teeth follows behind the captive for quite some time, refusing to relinquish 
its prize. Cubs seldom fight or quarrel. The young scampering around a 
prey trample large areas and bite off branches from tree and shrubs nearby. 
If prey is scarce in the locality of the young and their mother absent for 
a long time, hungry cubs will gnaw at rotten wood. 

Cubs remain with their mother for two, sometimes even three years, and 
begin to live independently only in their fourth year. This biological feature 
of tigers has been remarked upon before. "It has been said that old babry 
feed their cubs for three years, during which period the latter are so quiescent 
that they can be caught without fear of attack" (Rychkov, 1762). In 1953 
on Sikhote-Alin, I.T. Trofimova, the tiger catcher, came across a family 
of three three-year-olds and caught them all without much difficulty. 

By two years of age young tigers weigh over 100 kg. In the winter 
of 1950/1951 in the Ussuri region '4onchakV' (in their second year) [i.e., 
yearlings] weighing 113 kg each were caught (K.G. Abramov). A three- 
year-old male sometimes weighs up to 150 kg. Young tiger of this size 


generally hunt with their mother. From time to time cubs from two different 
litters have been sighted with a tigress, one or two being cubs from an earlier 
147 Males and females usually attain sexual maturity by their fourth year. 

In the Moscow Zoological Garden three female Ussurian tigers attained 
sexual maturity between the ages of three years four months and three years 
eight months. One male was sexually mature at the age of three years eight 
months; he attempted mating, however, at the age of two years seven months 
(Afonskaya and Krumina, 1956). Since cubs remain with their mother for a 
long time, she only litters once in two to four years. It is known that ' 'babry 
. . . whelp once in three years" (Rychkov, 1762). In the Moscow Zoological 
Garden, however, one female whelped twice in the same year and became 
pregnant a third time (Salmin, 1940). This female did not suckle her cubs. 

During the course of her lifetime a tigress generally produces 20 to 
30 cubs of which usually one-half perish within six months (Berten, 1954'''). 
In the Soviet Union a tigress produces only 10 to 15 cubs (A.A. Sludskii). 

According to observations made in captivity a tigress retains her ability 
to reproduce up to 20 years of age and her longevity is 40 to 50 years 
(Baikov, 1925). The Ussurian tigress Sirotka bom in the Moscow Zoological 
Garden lived for only 18 years. In India one tiger survived in a relatively 
small region for 15 years but was killed in its prime. Another predator 
attacked cattle of a particular region for 20 years and, when killed, was 
not at all senile (Pocock, 1939). 

Tigers molt twice a year; in the Ussuri territory molt takes place in 
September and October and again in March and April. The long and sharp 
claws are shed annually in late autumn. At the time of shedding their claws, 
tigers often scratch the soft bark of cedar, Amur cork, and fir. 

Enemies, diseases, parasites, mortality, ajid competitors. The adult tiger 
has no enemy in the animal world. Very rare indeed are instances of its 
death due to injuries inflicted by a male boar, buffalo, or bear. Tiger 
mortalities due to wild boar have been recorded in the Trans-Caucasus and 
Iran (Brandt, 1856), Kazakhstan, Middle Asia, the Far East and India. In 
the upper reaches of the Iman in the Ussuri region a wild boar gored a 
tigress to death, and her two surviving cubs died of starvation. In 1933 
in the Baikin region a tiger weighing 120 kg was killed by a boar. Near 
the Bol'shoi Sinanche River in 1913 a large brown bear crushed a tiger 
(V. Abramov, 1962). 

Cubs are killed more often than adult tigers. They are killed by male 
tigers, brown bears, and other predators. This was especially apparent in 
1956 when the yield of pine nuts and acorns in Birobidzhan was inadequate 

*Not in Literature Cited — Sci. Ed. 


and wild boar could hardly find food. Many brown bears wandered late 
in autumn because of the scarcity of food, instead of remaining in their 
lairs. One such bear on the Shukhi-Pokto range in the Khingan taiga attacked 
a three-year-old tiger and tore it to bits. Judging from the marks on the 
snow, the sequence of events may have been as follows. A tigress moving 
with a large cub attacked a herd of wild boar and wounded a pig. Leaving 
her cub with the quarry, the mother went away. A roaming bear came across 
the young tiger and the prey and a fight ensued between the predators. Tufts 
of fur from both the animals and numerous blood splotches were found 
scattered on the snow. The bear won. Having carried the pig to a wind- 
felled tree, the victor ate from it several times. After the boar was consumed, 
the bear returned to the tiger but was frightened off it by people (bCrivopusk, 
1957). Another instance of a wandering bear killing an adult tigress in 
Sikhote-Alin has been described above. In 1960 a young tiger was killed 
in combat with a bear in the Sikhote-Alin preserve (V. Abramov, 1962). 

In India tigers suffer greatly and sometimes even die from porcupine 
quills, which stick in their body when the rodent is attacked. A man-eater that 
was shot had 102 porcupine quills extracted from it. Some of these quills 
were 20 cm long and as thick as a pencil. Most were embedded in the muscles 
and some firmly lodged between bones (Corbett, 1957). In another dead 
adult tiger the liver and lungs were pierced at many places by porcupine 
quills (Buri'er, 1955)*. Evidently tigers in the Trans-Caucasus, Middle Asia, 
and southern Kazakhstan also suffer injuries from this rodent. 

The diseases suffered by tiger in the wild have not been studied. In 
148 zoological gardens they suffer from all infectious diseases characteristic 
of cats as well as those of other species of carnivorous animals. 

In one menagerie a Ussurian tiger suffered from food poisoning and 
died of it on the thirteenth day. The infection was transmitted by consumption 
of the flesh of a diseased horse (Vyshelesskii, 1948). Tigers also die of 
pasteurellosis, paratyphoid, carnivore distemper, etc. Two tigers caught in 
southwestern Tadzhikistan harbored five to seven tapeworms {Taenia 
bubesei) in the small and large intestines; these worms had formerly been 
recovered from African lion (Chernyshev, 1953; in the past the ranges of 
tiger and lion often overlapped). Toxocara mystax has also been detected 
in this predator (Mozgovoi, 1953). Toxocariasis is often noticed among 
tigers in zoological gardens. In an Amur tiger the following four species 
of helminths were found: Paragonismus westermanni, Physaloptera 
praeputiale, Dirofilaria ursi, and Uiteinarta sp. (V. Abramov, 1962). From 
a female caught in Tadzhikistan on June 26, 339 ticks belonging to four 
species were collected: Rhipicephalus turanicus (greater part), Boophilus 

*Not in Literature Cited — Sci. Ed. 


calcaratus (three), Hyalomma detritum (seven), and H. marginata 
(Chemyshev, 1958). Primor'e tigers are parasitized by the tick Dermacentor 
silvarum (Pomerantsev, 1946). 

The wolf is a competitor of tiger everywhere. It is known that at places 
in the Primor'e where tigers live, wolves are absent or very few in number; 
it is thought that tigers destroy wolves (Kaplanov, 1948; G.F. Bromlei). From 
the diligence with which a tiger chases dogs, one may presume that it will 
hunt wolf just as tenaciously. Leopard could also be a competitor of tiger; 
usually these two cats do not live in proximity to each other. Sometimes 
bear and yellow-throated marten trail behind a tiger to scavenge carcasses; 
jackal, jungle cat, and hyaena similarly follow tigers in Middle Asia. In the 
Far East, in the Amur valley, a male wapiti killed by a tiger was found by a 

149 Fig. 83. Track of a three-year-old tiger (male) in fine snow. Sankhobe River basin, 

Sikhote-Alin preserve. February-, 1960. Photograph by G.M. Veinger. 


brown bear who buried it in the snow (K.G. Abramov). In the Khabarovsk 
region in years of food scarcity (pine nuts and acorns) bear have no chance 
to fatten and do not remain long in their lair. Instead they set out to hunt 
more often than usual for wild boar and moose, thus competing with tiger. 
Under such circumstances bears sometimes chase young tigers off their 
prey (V.P. Sysoev). 

Crows and ravens invariably gather around the remains of a tiger kill. 
Small predators are not afraid of a dead tiger; in 1950 in Tigrovaya Balka 
preserve (Tadzhikistan) a tiger carcass was found which had been gnawed 
at by jackals. 

Population dynamics. It was mentioned earlier that the tiger population 
in the last century decreased everywhere and at present it has disappeared 
altogether in many parts of its range (Uzbekistan, Kazakhstan) or is only 
149 found as an intruder (Traas-Caucasus, Turkmenia, Tadzhikistan, southeastern 
Trans-Baikal, and Amur district). The reason for the rapid decline in the 
population of tiger was persecution by man and especially the indirect effects 
of his intervention (felling and burning of forests including the tugais, 
drainage, cultivation, and burning of reed floodplains, reduction in population 
of wild ungulates, and other activities). The main reason for the death of 
the last of the tigers in Kazakhstan was the extensive burning of reeds in 
the lower reaches of rivers during April and May every year, which destroyed 
all vegetation there in the 1930's through the 1940's (Sludskii, 1950). Tigers 
disappeared from the lower reaches of the Amu-Darya mainly because of 
fires (Pokrovskii, 1951). In Tadzhikistan their population decreased as a 
result of rapid cultivation of river floodplains (for example, Vakhsh). 

When the agriculture of Middle Asia and southern Kazakhstan suffered 
great losses as a resuh of wars (most of the large irrigated fields and cattle 
were destroyed) in the thirteenth and eighteenth centuries, the tiger population 
rose significantly. A similar phenomenon has been observed in China and 
India. Following major social upheavals (wars, revolts, and revolutions), 
which disturbed the economy of these countries, the tiger population began 
to rise. For example, in China the number of tigers rose significantly in 
the years of the National Liberation War which ended in 1949. 

Along with a gradual reduction in number of predators, temporary 
changes have also been detected in their population due to migrations, 
and mortality during unfavorable conditions, mainly scarcity or difficulty 
of access to food. In Sikhote-Alin in years of abundant yield of pine 
nuts and acorns, the wild boar population increases, and, subsequently, 
the number of tiger. The latter migrate there from northeastern China 
(Manchuria) and the Korean Peninsula. In the Amur district after the 
large-scale mortality of wild boar in 1955, tiger disappearance was almost 
complete (G. Modin). 


150 Fig. 84. Tiger tracks on wet coastal sand. Ta-Chingou area, Sudzukhin preserve. 

May, 1965. Photograph by V.E. Prisyazhnyuk. 

In the lower Amu-Darya a perceptible reduction in tiger numbers took 
place after an epidemic plague broke out among wild boar in 1938, deci- 
mating 50% of their population. After the emigration of wild boar from 
Amu-Darya delta in 1945, tigers disappeared from there almost completely. 
In some years, a perceptible rise in tiger population has been noted in the 
tugais on the right bank of the Pyandzh. Such phenomena occurred in 1936 
and 1950. In the latter year local hunters killed four tigers there in three 
or four months. The observed increase in tiger population on the Pyandzh 's 
right bank was associated with fires in the tugais on the left bank (in 
150 Afghanistan; Chemyshev, 1958). The tiger population may drop after winters 


151 Fig. 85. Track of a tigress on shoulder of a forest road. Upper Tudo-Vaku River 

in the hiian basin. Sikliote-AJin. September, 1967. Pliotograph by E.N. Mat>'iishidn. 

of abundant snow and snow crust prolonged into early spring. It is known 
that in such winters adult tigers suffer intense hunger and even die of 
emaciation (see above), but the young suffer even more since the mother 
cannot provide adequate food. Tiger mortality may possibly result from 
epizootic diseases breaking out among feline and other carnivorous animals. 

The main factors influencing tiger populations, apart from man and 
his activities, are climatic conditions which affect the population of ungulates 
serving as their prey, ease of access to food, and possibly epizootic diseases. 

Field characteristics. When moving fast a tiger appears to be a mono- 
chromatic light brown. It usually paces, striding almost step by step (Figs. 
82, 83, 84, and 85). The stride is 50 cm to 80 cm long or even 100 cm 
in particularly large males. In galloping, jumps measure 4.0 to 5.0 m in 
length and sometimes more. If the track had a length^^ of 20 cm or more, 

^*rhe length of a track is measured from the front point of the impression of the 
longest digit to the rear point of the sole, i.e., the rear of the padded portion (hunter's 
terminology). Invariably the size of the print of a front paw is given, since the forepaws 
are larger than the hind ones. 


the animal is large; that of a male is usually 16 cm x 14 cm and of a female 
15 cm X 11 or 12 cm. Depending on the width of the "heel" impression 
and other characteristics, four categories of adult and sex groups can be 
roughly identified: 1) "heel" width 10 to 15 cm — aduU male weighing 200 
152 kg or more; 2) "heel" width 9 to 12 cm — adult tigress weighing 150 kg 
or more; 3) "heel" width 6 to 8 and 9 cm — cub up to three years of age 
weighing 70 to 120 kg, in rare cases up to 150 kg; and 4) "heel" width 
3.0 to 5.0 cm — cub weighing 30 to 60 kg (V.K. Abramov, 1961). 

The tracks of a tigress are not so rounded as those of a male and the 
digital impressions are thinner (Fig. 87). The thick middle digits are 
particularly prominent in the male. Moreover, the sex of the animal can be 
readily determined from urine marks. The female urinates on the ground, 

151 Fig. 86. Track of a tigress willi cub. Old cut-over at liead-waters of tlie Ulaicha, 

Sikliote-Alin. December, 1969. Pliotograph by A.G. Pankiat'ev. 



152 Fig. 87. Schematic prints of front and left hind paws of a tigress; measurements 

149 mm x 154 mm and 131 mm and 150 mm. Ta-Chindzhan mountain. Sudzukhin 
preseive. Spring of 1947. Sketch by G.F. Bromlei. 

as a result of which a thawed patch is visible in the snow, while the male 
sprays urine on any prominent object such as logs, mounds, etc. Young 
males also leave a fairly prominent print but its depression is not great 
because of their lesser weight. An adult tiger's pace is fully equal to a man's 
stride while that of a young cub is shorter. When a three-year-old cub follows 
its mother, the tracks almost overlap; when mother and cub move 
independently two sets of prints are distinguishable. The presence of younger 
cubs is evident from multiple prints in a small area because they are playful. 
The well-being of aduh animals can also be judged from the shaфness of 
paw imprints. In the case of well-fed tiger the pug marks are sharp and 
deep; contrarily, in an emaciated one they are flat and shallow and wrinkles 
in the pads discernible (Fig. 87). 

Numerous fecal deposits are a sure sign of imminent approach to a 
site where the predator has consumed a prey. Feces are black or brown 
in color and undigested remains of the prey distinguishable — fur, bone, 
etc.; they are generally cylindrical, about 4.0 cm in diameter, and slightly 
pointed at the rear end. But when meat and blood predominate in the food, 
the feces are semiliquid, shapeless, and blackish. Both types of feces exude 
a sharp, putrid, long-lasting odor. That the tiger has fed in this area is 
confirmed by the discovery within a short distance of scattered vertebral 
bones, long bones and sometimes even the skull of a large ungulate. 

The roar of tigers is heard rarely even at places where they are quite 
common. Only during rut is the animal's roar common. Usually the first 
roar is protracted, repeated in quick succession, and terminates in three or 


153 Fig. 88. Remains of three-year-old tiger cub killed and eaten by a brown bear. 

Shukhi-Pokto preserve, Khabarovsk region. 1956. Sketch by N.N. Kondakov from 
a photograph by Krivopusk. 

four short roars. Sometimes a low throaty "ah-oun" or "ee-oh-unnu" is 
emitted. When enraged the animal issues a hollow snarl and produces 
coughlike sounds with an intense exhalation of air through an open mouth 
(Baikov, 1925). Moreover, tigers sometimes produce a high pitched sound 
which could be described as "muuk" or "puuk" and is somewhat similar 
to the bugle of a wapiti in rut. Some hunters and even zoologists assume 
that tigers entice wapiti through such a sound. In India at places where there 
are many sambars tigers imitate their call to attract them into ambush (this 
153 opinion requires confirmation). A contented and resting tiger rumbles* a 
long time but, on being frightened, issues a characteristic "vuuf". Tigers 
generally attack their prey silently. (A.S.) 

Practical Significance 

When tigers were numerous they caused some damage to animal husbandry. 
For example, in 1905 a pair of tigers living on the Karatal River (southern 
Pri -Balkhash) killed about 30 camels and 100 horses (Shnitnikov, 1936). 
As late as 1929, in the lower reaches of the Hi, tigers destroyed over 200 

*Myrlychet in Russian original; probably a misprint for murlychet — Sci. Ed. 


head of cattle (N. Sh., 1930). In 1938 a large tiger in Parkhar (Tadzhikistan) 
destroyed in one month 23 large and small cattle on a collective farm. Cattle 
particularly suffer from this predator in wintering sites located among reeds 
by rivers. Tigers inhabiting the Chu River attack wintering cattle, mainly 
cows, and more rarely camels (Gem, 1891). 

With the arrival of Russians in Middle Asia, i.e., from the middle of 
the nineteenth century, intense hunting of tigers began. Moreover, their 
destruction was encouraged. In the 1870's a prize of 25 rubles was awarded 
for every tiger killed and at the beginning of the present century raised 
to 50 rubles. Even as late as 1929 in Semirech'e a special decree was issued 
for killing tigers. Tigers could be killed by any method and a bounty of 
100 rubles collected. All these measures and some other factors led to the 
rapid extinction of the predator. In the 1930's and later in Middle Asia 
tigers continued to attack cattle from time to time only in Tadzhikistan and 
the Amu-Darya delta, but this loss was extremely small. Formerly, tigers 
sometimes harmed cattle even in the Far East. In June, 1869 at Troitsk, 
on the shore of Lake Khanka, tigers killed in one month 32 cows and horses 
(Przheval'skii, 1870). In the last decade the damage caused by tigers to 
cattle has been negligible even in these regions. 

At present the damage done to game husbandry by tigers is negligible 
because of their small population. Moreover, zoologists who have studied 
tigers in the Far East in the last decade consider them useful to game 
154 husbandry, since wolves are absent or few in number at places where tigers 
are present. Tigers do kill a few ungulates but, unlike wolves do not drive 
them away (Salmin, 1940; Kaplanov, 1948; G.F. Bromlei). 

Several instances of attacks on man by tigers have been described in 
the Soviet Union in the last century. They occurred mainly in Middle Asia 
(excluding Turkmenia), Kazakhstan, and the Far East. Usually animals shot 
and wounded or chased by hunters attacked, and only very rarely did an 
attack occur without provocation. Nevertheless, in the lower reaches of the 
Syr-Darya a tiger reportedly attacked and tore to pieces a woman collecting 
firewood, and an unarmed military officer passing in June through reed 
thickets ("Vernenskii Grazhdanin," 1880). Instances of tiger attacks on 
shepherds have been reported from the lower reaches of the Hi (A.M. 
Nikol'skii, 1885). Attacks on man in the Far East were recorded in the middle 
and third quarter of the last century. In 1867 on the Tsymukha River, tigers 
mutilated and killed 21 men and wounded 6 others (Przheval'skii, 1870). 
In the present centur}' no tiger attacks on man without provocation have 
been reported (Kaplanov, 1948; Abramov, 1960; G.F. Bromlei). In preserves 
where tigers are conserved today they pose no threat to man because they 
are not hunted here and wild ungulates are bountiful. 

Tnie man-eaters, i.e., animals for which man is a regular, if not the only 


155 Fig. 89. Young tiger pressed to the ground with a "fork". Photograph by V.G. 


prey, are now absent in the Soviet Union. This is due to their relatively 
small population and the abundant availability of ungulates at places where 
they live. In India, however, animals regularly attacking man were numerous 
in the past and some still exist today. In the middle of the last century man- 
eaters were a common phenomenon in some regions of central India. In 
the Monula region alone they killed 200 to 300 humans in one year and 
decimated entire villages (Jerdon, 1874). Even in 1924 tigers killed 603 
humans in British India (K-li, 1926). One tigress in Nepal killed 200 
humans and in Kumaon over a four-year period before she was herself 
destroyed — 234 (Anderson, 1964). In the central regions of India about 


50 men were killed by tigers every year in the last decade (Berten, 1954*). 

Man-eaters are usually animals compelled to feed on humans, an un- 
natural food for them, being unable to hunt healthy, strong animals. The 
reason for such behavior in nine out of ten cases is a wound, and in the 
remaining case old age (Corbett, 1957). The wound might be the result 
of an unsuccessful shot by a hunter or the predator's attack on a porcupine. 

In China attacks on woodcutters and coachmen have been reported 
in Kirin province (Baikov, 1925). Instances of tigers entering cabins and 
carrying off adults and children are also known. Once a tiger attacked and 
killed three farmers in a forest but did not feed on the carcasses. In the 
southern part of China tigers killed 60 men in a short period of time (Allen, 
1938). Attacks on humans by tigers have occurred in recent years in Yunnan 
province, even as late as 1949 (A.A. Sludskii). 

Tigers are valuable fur-yielding animals but the importance of their 
skin as an item of clothing is negligible today. Throughout the Soviet Union, 
before hunting was banned, only ten animals were caught annually and their 
skins usually retained by the people; three or four were killed in Tadzhikistan 
and five to eight in the Far East. Only one to four skins were received for 
curing (G.F. Bromlei). Formerly, most skins were retained by the hunters 
or bought by collectors. Only three tiger skins (from Kirgizia steppe) passed 
through the custom houses at Petropavlovsk, Presnogor'kovsk, Om'sk, and 
Korkovsk from 1857 to 1861 even though dozens of tigers were killed 
annually in the nineteenth century (Krasovskii, 1868). 

The number of tiger skins coming from the USSR in relation to the 
world output is almost negligible. The world output from 1807 through 1910 
was estimated as 500 skins (Kaplin, Ivanov, and Patushenko, 1955); Russian 
fairs annually received only 30 to 50 skins up to 1914. In the 1920's around 
60 tigers were caught in the Soviet Union, including two or three live ones 
155 (Solov'ev, 1926). In these same years the annual catch in Amur and Primor'e 
provinces was 25, in Korea 25, and in northeastern China 50 to 60 (Baikov, 
1925). Amur tiger skins are especially prized in the world market. 

The rising price of tiger skins is one of the factors responsible for their 
persecution. For example, in 1760 a large tiger skin in Orenburg fetched 
13 rubles; a medium-sized one 8 mbles, and a small one 5 rubles (Rychkov, 
1762); in the early 1900's the price exceeded 100 rubles (Z^rudnyi, 1915) 
and even rose to 200 rubles in Middle Asia. The skins of Far East tigers 
were even more expensive. In the Primor'e region a tiger skin cost 70 rubles 
in 1886 and as much as 100 to 200 rubles in 1895 (Silant'ev, 1898); at the 
same time sable skins sold for 5 to 25 rubles each. In 1927 and 1928 in 
the Far East a tiger skin was valued at 300 to 500 Mexican dollars (Baikov, 

*Not in Literature Cited — Sci. Ed. 


1928). The present market price of a cured tiger skin varies from 4 to 35 
mbles depending on its quality. 

The carcass of a tiger is no less valuable than the skin; even in the 
1930's tiger carcasses were exported to China, where they were sold to 
pharmacists. The people of China, Semirech'e, and the Kul'dzhinsk region 
eagerly bought the heart and bones of tiger (Gem, 1891). In 1929 and 1930 
in the Far East tiger carcasses fetched 3 rubles 50 kopecks per kg, and the 
bones 15 rubles per kg. During this same period in northeastern China, 
10 Mexican dollars were paid for 16 kg of carcass (Baikov, 1928). A hunter 
could thus receive up to 1,000 rubles for a single adult tiger. 

Formerly in the Far East, caught animals were not skinned but were frozen 
whole and dispatched to the nearest town where they were bought by Chinese 
marketeers and transported to interior China, mainly Peking and Tientsin. 

Chinese medical practitioners highly prized the carcass of an adult male 
and its various parts. Medicines were prepared from each organ which cured 
not only different diseases, but also cowardice, emaciation, etc., and talismans 
were made from them. In the middle of the last century Kazakhs generally 
cut out the claws of a tiger and stitched them on wearing apparel for children; 
claws were considered charms which drove evil spirits away from a child 
(Karelin, 1847). 

Even in 1958 tiger bones were sold in China. A single animal yielded 
sometimes up to 30 kg of bones. Particularly prized were the bones of the 
leg, especially the patellas (Sludslcii, 1966). Various body parts of the tiger 
have been used in Korean medicine. Both Chinese and Koreans enjoy tiger 

From very early times tigers have been caught live and exhibited in 

circuses or kept in menageries. Roman emperors often displayed their tigers. 

156 In the Middle Ages tigers were trained in the east for hunting big game 

(Simashko, 1851). From 1260 to 1280 Kublai Khan, grandson of Genghis 

Khan, regularly used tigers on hunts in China (Marco Polo, 1955). 

In ancient times in the Trans-Caucasus, Middle Asia, and Kazakhstan, 
tigers were hunted for sport and for the purpose of their destmction. In 
Azerbaidzhan, Armenia, and Georgia, czars and aristocracy used to arrange 
hunting parties for tigers. Tiger hunting was very prevalent during the period 
of Mongol mle. Mongolian aristocracy and military detachments hunted tigers 
in order to perfect military techniques and to inculcate bravery in the soldiers. 
In the Middle Ages Mongolians and peoples elsewhere regarded the tiger 
as a superior being and were afraid of it (Grekov and Yakubovskii, 1950). 

With the decline of the Great Khans, hunting of tigers also declined, 
but with the arrival of Russians in Middle Asia in the nineteenth century, 
it was revived on a large scale. Soldiers from border regions, special ' 'hunting 
units" of the army, and individual hunters, mainly army officers, began 


hunting tigers. Tracking and catching tigers (and wild boars) helped instill 
bravery in the Kazakhs and soldiers. Regular hunting of wild boars and 
tigers on the Syr-Darya greatly influenced the qualities of courage of local 
garrisons, and inculcated valor and military dexterity in the soldiers who 
"became fearless because accustomed to facing danger, and later such 
garrisons distinguished themselves in many notable battles in the Syr- 
Darya region" ("Vernenskii Grazhdanin", 1880). 

The Kazakhs of Syr- Darya and Amu-Darya garrisons hunted tigers by 
lying in wait with their guns or by shooting animals beaten out of rushes. 
The hunt was sometimes conducted on horseback or sometimes small groups 
charged as in a bayonet attack. Almost every such attack resulted in the 
death or wounding of soldiers and officers. 

The local people of Middle Asia hunted tigers by attaching a knife 
to a "spring" on a track or at the site of a killed calf. A flexible pole with 
the knife tied to it was lowered to the ground and fixed with a simka^^ 
in such a way that the animal on touching the latter released the pole. As 
the pole straightened the knife ripped the animal's underbelly. In the Amu- 
Darya a large automatic type of trap used by northem trappers was employed. 
In the tugai a strap tug supported a tiny pole fitted with sharply pointed 
knives. A careless tiger on touching the strap released the tug and the pole 
with the knives hit the flanks of the animal. Kazakhs from time to time 
set up log mats with nails and knives on the outer logs. All these methods 
were usually ineffective and the animal often escaped with wounds; 
nevertheless one or two were caught in a season (A.I., 1888). More 
commonly tigers were caught in large portable traps and shot. 

The most prevalent method of hunting tigers was to set up around the 
remains of its kill one or several guns which went off when the animal 
touched a string. As in the previously described methods of hunting, the 
powerful animal, though shot, was rarely killed on the spot. Injured, it fled 
and later became dangerous to man. 

In the last century Kazakhs sometimes caught tigers by poisoning the 
remains of its prey with a poison prepared from the roots of Eminiiim 
lehmanni. Sometimes hunters lay in wait for the animal at the site of its 
kill or on its path after constructing a wooden platform protected from all 
sides. The platfoim was fitted on poles firmly driven into the ground. The 
tiger was shot from the platform and if it attacked, killed from above with 
a bayonet or large knife. Sometimes Kazakhs joined three such platforms 
together and firmly tied sticks on top. A thin strap was fastened to the 
top of this cage. Several men sat inside the cage equipped with guns and 
other shaф weapons. Another method was to go to the animal 's den wearing 

^^A fine string used in making various traps. 


157 the protective device described by Gem (1891). Kazakhs also hunted tigers 
in groups of 30 to 40 men mounted on horses, killing the beast with bow 
and arrows (Eversmann, 1850). 

In the Far East local hunters (Udekheits, Evenks, Crochets, and others) 
did not hunt tigers, as they regarded the species with reverence and caught 
one only under exceptional circumstances (see below). In the nineteenth 
century Russians bagged tigers with both bow and gun. A bait such as a 
slaughtered wild boar or roe deer was placed in the animal's path. More 
often a dog served as bait and loaded guns were set up around it (Kulagin, 
1923). The animal, wounded by the automatic guns, sometimes attacked 
the men chasing it. More rarely, the animal was poisoned with strychnine. 

Tiger skins were med in making carpets and rugs. 

In ancient Rome live tigers were imported from Persia and Armenia. 
In the eighteenth century animals from Bukhara were brought into the 
menageries of Russian czars (P.I. Rychkov, 1762). There were still tiger 
catchers in the Ussuri territory from 1902 to 1906. Some farmers caught 
live tigers with their bare hands, using neither cage nor trap, and tied the 
animals up with ropes (V.K. Arsen'ev, 1949). 

There was a particularly great demand for live Ussurian tigers in the 
1920's. At that time an adult animal fetched 1,000 pounds steriing (about 
10,000 gold rubles) and a cub of up to one year, 100 pounds. The demand 
for live tigers within the Soviet Union and abroad continues even now. 
A young Ussurian tiger in the 1950's to the 1960's fetched up to 500 rubles 
or more. On the international market a Ussurian tiger in the mid-1960's 
cost about $4,000 and a southern Chinese tiger about $3,000. The great 
demand for live tigers resulted at one time in the capture of almost every 
cub produced annually. 

Usually a group of four or five hunters on finding the tracks of a tigress 
with cubs pursue them, often for several days. On closing the gap the hunters 
yell and chase the animals, shooting in the air in order to separate the tigress 
from the cubs. This chase sometimes covers 5 to 10 km. The weaker of 
the cubs begins to lag behind its mother, who continues to move forward. 

158 Having lost sight of its mother, the frightened cub strays from her path, 
turning sideways. Dogs, kept leashed until that moment, are then set on 
its course. ' 

The released dogs frighten the cub until it seeks shelter under an 
upturned or wind-felled tree, or in rocks, etc. On reaching the cub and 
restraining the dogs, the hunters roll and pin the cub to the ground with 
poles with forks at the end. They then tie its legs with straps or rope loops 
and muzzle it with a pre-prepared muzzle (K.G. Abramov, 1956). 

This is a common method of catching tigers live, but quite often another 
is employed. Sometimes the cornered animal is poked by one or two men 



Fig. 90. Tied cubs. Primor'e region. Winter of 1952/1953. Photograph by N. Nazarov. 

with the butt of a rifle or a padded jacket or bag. Before it can spring for 
an attack other hunters overpower it and, holding it by the nape of the neck 
and paws, pin it to' the ground where it is trussed up (Shklyar, 1935; Reutov, 

Having caught the first cub of a litter, one of the hunters stands guard 
over it while the rest start chasing the remainder until the entire litter is 
caught. Rarely is a cub capable of eluding the hunters. A group of coastal 
tiger catchers detected 17 litters comprising 48 cubs from 1936 through 
1957 and caught 45 of them (V. Abramov, 1962). 

The efficiency of the methods described above for catching tigers can 
be judged from the following data. From 1930 through 1940 on the Iman 
and its tributaries alone, 40 cubs and even 9 adult tigers were caught (Sysoev, 
1955a). One tiger hunter caught 21 and killed 18 (up to 1935); another 
caught 36 in his lifetime, and still another caught 16 and killed 8. Catchers 
from Loumo from 1947 through 1955 caught 24 tigers of which some 
weighed 116 kg (cubs are caught live up to the age of three years inclusive, 
when they quite often weigh over 100 kg). 

In the remote past, among peoples of the Far East, and also China, 
Vietnam, Japan, Mongolia, Korea, India, Nepal, Indonesia and other 
countries, there existed a special tiger cult. Tigers and dragoas were depicted 


as objects of art in palaces of rulers, huts of cultivators, cabins of animal 
catchers, and on altars as idols. In the Altai, in excavations of the Pazyryk 
burial mounds, circa 500 to 300 B.C., some lifelike drawings of tigers were 
found together with animals with the body of a tiger or the head of a tiger 
on the body of an eagle or vulture (Rudenko, 1953). In the palace of Toprak 
Kal, in ancient Khorezm, tiger illustrations have been found (Tolstov, 1951). 
In the forests of the Far East animal catchers and those seeking ginseng 
raised idols in order to propitiate the tiger. 

Among the Chinese, Udekheits, and other peoples, tigers aroused a 
special mystic fear and reverence; the dark pattern on the head of a tiger 
resembles the hieroglyph "van-da," which means "great emperor" (Fig. 
91). According to popular belief, the spirit of a ruler lived in the body of 
a tiger as a means of expiating his sins. Formerly, Udekheits killed a tiger 
only if it attacked their camps. The tiger cult (mafa cult) among these peoples 
resembled the cult of the "master" of nature — of the taiga and the universe. 
Myths of a totemistic nature were associated with the tiger (Ivanov, Levin 
et al., 1956). The tiger cult prevailed even among the Crochets. They killed 
a tiger in accordance with the laws of their native land, only to avenge 
the killing of their kinsmen. Nanaiets also had a special reverence for the 
tiger. The members of one family traced their ancestry to this animal. All 
the foregoing beliefs reveal the important role tigers formerly played in 
the life of peoples of eastern Asia. 
159 As a gift of nature and history the tiger is of paramount scientific 

importance, which is increasing every year due to the animal's rapid decline. 
The tiger is no longer dangerous nor harmful to man in the Soviet Union. 
Every available measure will have to be implemented to prevent its total 
destruction. Such measures include a total ban on tiger hunting and severe 
restrictions on catching of cubs throughout Soviet territory. In the Khabarovsk 
and Primor'e regions all types of tiger catching were totally banned from 

158 Fig. 91. Hieroglyph "van-da" (right) meaning "great emperor" and dark -colored 

pattern resembling it on head of a tiger. Sketch by N.N. Kondakov. 


1947 through 1954. In 1954 a temporary relaxation was granted for catching 
cubs. In the Primor'e region in 1956 shooting and catching of tiger were 
banned throughout the year and throughout the territory for a period of 
five years. At the same time a penalty of 5,000 nibles (old currency) for 
unauthorized tiger killing was imposed. Hunting of tigers had been banned 
even earlier (1952) in this region. The ban on catching and killing of tigers 
in the Khabarovsk and Primor'e regions continues to the present day (1968). 
However, beginning in 1960 special licences have been issued for catching 
a limited number of tiger litters. 

Measures for protecting the Far East tiger have produced positive results. 
The decline in its population, counted at the beginning of the 1940's, has 
been arrested at about 120 to 140 individuals (see above). In Tadzhikistan 
the killing of tiger year-round was prohibited in 1957 but, sad to say, the 
ban came rather late. 

Tiger conservation requires an international agreement with governments 
of neighboring countries in which this animal still survives (Korean 
Democratic Republic, South Korea, China, Afghanistan, and Iran). It is 
necessary to set up large sanctuaries in the regions adjoining the Korean 
Democratic Republic, Chinese People's Republic, Afghanistan, and Iran, 
and to enlarge the Sikhote-Alin, Sudzukhin, and Suputin preserves (the upper 
Suputinka, originating in the Dadyan'-Shan range in the western spurs of 
Sikhote-Alin). A preserve is necessary in Birobidzhan for preserving tigers 
in the Shukhi-Pokto range, and game refuges should be set up in other places 
where tigers still survive and steps taken toward the conservation of ungulates 
in them, which serve as food for the predator. Raising tigers in zoological 
gardens would help to increase their number in the Soviet Union. 

The implementation of these and other measures would ensure the 
survival of this most interesting animal in our fauna; if unheeded, this cat 
will be totally extinct in a few years. 

Tigers are protected in several foreign countries. In the Chinese People's 
Republic the hunting of Amur tigers inhabiting the northeastern province 
is totally banned. Measures for the protection of tigers have also been effected 
in the Korean Democratic Republic. In India tigers are protected in preserves 
and in a series of long-term game refuges. In the Union of Burma tigers 
are likewise conserved in long-term game refuges. The Javan tiger* in 
Indonesia is protected in a sanctuary in the western part of Java. (A.S.) 

*Now extinct — Sci. Ed. 

Panthera pardus Linnaeus, 1758 

1758. Fells pardus. Linnaeus. Syst. Nat., 10th ed., p. 41. Egypt. 

1856. Felis tul liana. Valenciennes. C. R.Acad. Sciences, vol. 42, p. 1039. 
Ninfy, 40 km east of Smyrna (Izmir), Asia Minor. 

1857. Felis orientalis. Schlegel. Handl. d. Dierkunde [sic], vol. 1, p. 23. 
Korean Peninsula. 

160 1862. Leopardus japonensis. Gray. Pfoc. Zool. Soc. London, p. 262. 

Northern China (ascribed in original description to Japan where 

leopards are absent and were never present). 
1867. Felis fontanieri. Milne-Edwards. Ann. Sc. Nat. Zool., vol. 8, p. 375. 

Near Peking. 
1867. Leopardus chinensis. Gray. Proc. Zool. Soc. London. Mountains west 

of Peking. Non Felis chinensis Gray, 1837. 
1878. Felis leopardus. Sclater. Proc. Zool. Soc. London, p. 289. Persia (?). 

^"The correct name in the Russian language is "bars" and not leopard. Intense efforts 
were made in the last decade to introduce the latter name into Russian literature. The word 
"bare" has been in popular usage and literature for several centuries (at least from the sixteenth 
century) and right from the beginning it referred to the Caucasian leopard, i.e., io P. pardus. 
Among Russian peoples this name is used throughout the Caucasus. The word "panther" 
was used for the first time at the end of the nineteeth century and only in the territory of 
the Caucasian preserve among hunters; the word "panther" was introduced during the period 
of the great imperial hunts from Petersburg. Russian peoples called, and continue to call 
the animal "bars" throughout Turkmenia, in the Soviet Far East, and in northeastern China 
(Manchuria). This name was used in Russian zoological classics of the eighteenth, nineteenth, 
and early twentieth centuries (S.I. Ognev, for example), and is still used by many well- 
known contemporary writers. It has long been listed in Russian dictionaries with a reference 
to P. pardus (". . . like a desert bars, ferocious and wild ..." or "... the desert's permanent 
guest — mighty bars . . ." Lermontov; or "wearing a bars skin," etc.). 

In the distant past, in the eighteenth century, as can be seen from Pallas (1811), but 
evidently even before, the word "bars" was popularly used for irbis [ounce] (Uncia uncia) 
in Siberia — and later in Semirech'e, and Middle Asia. This is quite natural considering the 
similarity of these two species and such confusion is not uncommon. Furthermore, in the 
fur trade of the seventeenth century, references also occur to the skins of "irbizy". In the 
recent period (nineteeth to the early twentieth century) in Russian zoological literature the 
name "snow leopard" (identical in concept to the English, German, and French names) 
and "irbis" (ounce) (from Turkey and Mongolia) has been finnly established for Uucia 
uncia. The term "bars," as earlier, is thus retained for P. pardus. 

There is no sense in changing the name "irbis" to "bars" nor introducing the new 
name "leopard" for "bars"; neither is there any scientific justification for doing so. Neither 
is there zoological or philological justification, nor any practical convenience achieved. Such 
changes would only lead to misunderstandings, confuse nonspecialists altogether, and 
needlessly clutter up the language. (V.H.) 


1903. Felis villosa. Bonhote.A«/L Mag. Nat. Hist., vol. 11, p. 475. Amur 

1904. Fells pardus grayi. Trouessart. Cat. Mammaliwn, p. 268. New name 
for chinensis Gray. 

1914. Leopardiis pardus ciscaucasicus. Satunin. Сотр. Mamm. Imp. Ross., 

vol. 1, p. 159. OpredeliteVmlekop. Ross. Imp., vol. 1, p. 159. 

"Forested region of northwestern Caucasus (montane zone of the 

Kuban district.") 
1927. Panthera pardus saxicolor. Pocock. Ami. Mag. Nat. Hist., ser. 9, 

vol. 1, p. 213. Astrabad, Iran. 
1959. Panthera pardus dathei. Zukowsky. Zool. Garten, vol. 24, p. 348. 

Sandabad, southeast of Shiraz, southern Iran. 
1964. Panthera pardus transcaucasica. Zukowsky. Zool. Garteti, 28, 4, 158. 

"From Georgia to Armenia and Azerbaidzhan" {sic\ — V.H.). (V.H.) 


Color yellow and reddish-yellow with small black rosettes and spots with 
a well-defined light colored center. Body length less than 160 cm, 
condylobasal length of skull less than 250 mm, and length of upper camassial 
tooth less than 29 but more than 21 mm. (V.H.) 


Size large. Much smaller in size than the tiger or lion, the leopard, snow 
leopard (ounce), and cheetah are nonetheless bigger than the remaining 
Russian cats and represent the next largest species in the family (Fig. 92). 
Body long, with rather short legs; very agile, light and slender, with 
a long tail (length more than one-half length of body), and a relatively small 
rounded head. Forehead bulges, facial portion moderately long, and ears 
short, rounded, and wide-set. Eyes small, with round pupils. Mane or longer 
hair on dorsal side of neck and cheek absent; hair of very uniform length 
throughout the body, relatively short and close-fitting, and not luxuriant 
even in winter. Type of hair frequently responsible for slender appearance 
of animal. Body, however, very strong and muscular with powerful and 
broad front paws. 
161 "Pelage short, coarse, and sparse. There are about 3,000 hairs per ст- 

оп the back, with only four underfur hairs to one guard hair. A characteristic 
difference is apparent in the length and thickness of black (in the spots) 
and yellow hair; the former (as in all fur-bearing animals) is thinner and 
longer. Yellow guard hairs and 30 mm long and 121 microns thick but the 
black hairs 40 mm long and 96 microns thick. The length of yellow top 



^W г 



161 Fig. 92. Far East leopard, Panthera pardus orientalis Schleg. Vil'nyus Zoological 

Garden. Photograph by B.B. Marni. 

hair (four categories) is 26, 25, 24, and 22 mm, and thickness 109, 92, 56 
and 37 microns; the corresponding measurements for black hairs are 30, 
31, 29, 27 mm, and 89, 64, 57, and 49 microns; the length of yellow underfur 
is 20 mm and thickness 21 microns and of black underfur 24 mm and 32 
microns" (B.F. Tserevitinov; specimen from Middle Asia). Differences 
between the summer and winter coats vary in different subspecies but 
generally small. In Amur leopards the length of hair on the back in summer 
is 20 to 25 mm and in winter 50 mm. 

The color of leopard is distinguished by a general light background 
of yellow or reddish-yellow color with black spots of two types — solid 
or annular (rosettes). In the center of the latter a light-colored field occurs, 
corresponding fairly well in color to the main background of the coat. The 
black ring may be complete but is usually interrupted at two to five places, 
i.e., it consists of two to five separate spots grouped into one ring. Transverse 
(vertical) stripes are not present but sometimes individual spots on the back 
fiLse into short longitudinal stripes. The outlines of the spots are well defined. 

The general background color varies individually, geographically, and 


seasonally. Among Soviet leopards it changes from a fairly dull light yellow 
or yellowish-gray to a bright yellowish-red with a golden tinge, but may 
also be light gray or almost white or whitish. The color is most vivid on 
the back, and often on the dorsal part of the neck and head. It gradually 
lightens and loses the yellow and red tinges on the flanks, and is white 
on the abdomen, inner side of the limbs, the throat, below the neck, extremity 
of the legs, the underside, and the terminal half and upper side of the tail. 
The color contrast between the back and rest of the body is variable and 
depends on the general intensity of coloration. This contrast is less sharp 
in light-colored forms than in reddish-colored animals. 

The extent and character of spottiness is variable. Spots are usually pure 
black in color but may also be brownish. The total number and density of 
162 disposition of spots is likewise variable. Also variable are the size and form 
of solid spots and size and structure of annular spots, i.e., the number of 
their constituent elements, their rounded or oval form, relative numbers 
in the general pattern of skin and disposition on the body, and color of 
the central field (corresponding to the yellow background of the skin or 
darker), etc. The anterior part of the muzzle is devoid of spots and small 
dark markings occur only in the region of the vibrissae and a dark spot 
on the lower lip in the comer of the mouth. On the cheeks, forehead, between 
the eyes and ears, and along the top and sides of the neck, solid black spots 
of various sizes, but generally small, are present; on the dorsal side of the 
neck the spots are somewhat elongated. Back of the ears black. 

Annular spots occur on the back and flanks. Spots on the spine are 
usually elongated or form a pattern of elongated-annular and large elongated 
solid spots. Sometimes individual spots fuse into a short longitudinal stripe. 
Annular spots, consisting usually of a few sections and measuring up to 
50 to 65 mm in diameter, are distributed throughout the flanks, on the top 
of the shoulders and on the thighs or in their upper parts. While the annular 
spots on the flanks descend to the abdomen and usually comprise the 
predominant type of spots (some are distributed lower), their distribution 
on the shoulders and thighs is more variable. Thus on the shoulder and 
forelegs they may be present together with solid spots and even descend 
to the forearm or even be absent on shoulder. They may also cover the 
entire region of the thighs or only their upper part. Solid spots occur all 
over the body, wherever annular spots are absent, but sometimes are present 
together with several or many annular spots. They are few on the flanks, 
close to the spine, and more numerous toward the abdomen. The legs and 
abdomen are covered with solid spots. The spots on the outer side of the 
legs are large toward the top and smaller down below, becoming tiny freckles 
on the wrist and paw. 

The top of the tail and part of its underside are covered with large annular 


or solid spots. The vibrissae consist of black, white, and half-black, half- 
white resilient hair up to 110 mm long. The claws are a light waxy color, 
highly compressed from the sides, steeply curved, and very sharp; their 
length along the outer arc reaches up to 55 mm (front paw). Claws on the 
hind paws are smaller and not so sharp. 

Seasonal color variability and the nature of the coat in Soviet leopards 
are distinctly manifested but not to the same degree in all forms. In spite 
of long hair, up to 50 mm in winter, the pattern of spots is shaфly maintained 
as in summer. The main background color in winter is paler and duller 
than in summer, but quite vivid even in winter in brightly colored forms. 
In pale forms it may even be whitish, resembling the color of the ounce. 

Individual color variability is generally confined to changes in vividness 
of color of the main background and the pattern of spots (see above). Two 
strikingly different color types — duller or more vivid — are seen in some 
populations. These are not connected, or almost not connected, by inter- 
mediates (Rerov, 1935; Tsukovskii, 1959; also see ''Geographic 
Variation").^^ Color macromutations affecting spottiness, described from 
India (Pocock, 1941), are not known among Soviet leopards. Melanistic 
animals ("black panther") are quite common in some parts of southern 
Asia (Sikkim and Nepal — P. p. pernigra Hodg., and Java — P. p. melas 
G. Cuv.), but absent or extremely rare within the Soviet Union.^"^ Young 
leopards have a very light background color, which is yellowish-white on 
the back and whitish on the flanks. Their spots are brown or brownish- 
chestnut, similar to the color of the back of the ear. Spottiness is very dense, 
however, and hence the animal appears quite dark. 
163 Geographic color variation pertains to shade of color and vividness 

of the general background (from a very light grayish-white or pale yellow 
to a bright and full reddish-chestnut) and in several details of the nature 
of the spot pattern (see "Geographic Variation"). 

The skull of a leopard differs from that of a tiger in many features 
(see description of tiger), primarily size. Even in very old animals the skull 
exhibits less prominent "predatory" features and is more "catlike" (see 
characteristics of the family). In leopard the interorbital and postorbital 
coastrictions are smaller, the cranium larger, zygomatic arches less separated, 
facial part relatively less developed, etc. 

^■^hese, however, do not represent a dual-phase color but extreme types of variations. 
The absence or rarity of animals with intemiediate colors is explained by the nonavailability 
of a large series of skins in museums. 

^One report (Flerov, 1935) is available pertaining to southern Tadzhikistan, but quite 
possibly it could also pertain to the snow leopard. 

The absence of melanistic animals in the Middle Asian and Caucasian parts of the 
geographic range, which are known for their aridity, particularly the former, is quite expected. 


The skull of leopards is characterized by the following features. On 
the whole it is relatively massive, fairly long with rather close-set zygomatic 
arches, and quite low. Upper line of profile from highest point in interorbital 
region to rear runs relatively gently, but forms a very steep arc toward the 
front. Frontal area almost flat or with faint longitudinal depression. Crests, 
especially sagittal and lambdoidal, highly developed (Fig. 93). 

Rostral width (above canines) in adult animals wider than postorbital 
constriction and more or less equal to interorbital width. Zygomatic arches 
strong and zygomatic process of temporal reaches base of postorbital process 
of jugal. Postorbital process of jugal does not reach level of lacrimal foramen. 
Lower anterior region of orbits slightly thickened and bluntly rounded toward 
the top. Infraorbital foramen fairly large, its maximum diameter exceeds 
width of bony part between it and edge of orbit. Nasals long and fairly 
uniformly narrowed in posterior part. 

Hard palate relatively narrow and extended; its width much shorter 
(by 35 to 50%) than its length. Posterior edge of bony palate in the middle 
of inteфterygoid vacuity (along the suture between the palatines) with small 
pointed projection. 

Tympanic bulla (see Fig. 23) relatively large, greatly inflated, with a fairiy 
smooth surface. Anterior chamber (p. ectotympanica) small and occupies 
much less than one-half of volume of entire bulla. Its boundary (usually 
poorly visible) is shifted in an anterolateral direction. Anterior margin of 
bulla lags far behind glenoid fossa and postglenoid process, by not less 
than one-half width of interpterygoid vacuity. Distance between inner walls 
of tympanic bullae more than width of interpterygoid vacuity. Jugular 
foramen large. Paroccipital process massive and closely adheres throughout 
to posterior wall of tympanic bulla and does not form a free falcate process. 

Canines relatively thin at base but long and shaф. Additional fifth 
tubercle absent on anterior outer part of upper carnassial tooth. 

Sexual dimorphism reflected in small overall size and slender skull 
stmcture of female. In males, as a result of the far more intense development 
of masseter muscles, the entire cranium is more compressed. Frontal part 
of cranium and postorbital constriction ("waist") long and highly 
compressed; cross section ("postorbital width") roughly equal to inter- 
orbital width. This isthmus in females is usually short and the compression 
there less: width of this region more than interorbital width. Crests, especially 
sagittal and lambdoidal, less developed and teeth on the whole somewhat 
weak; canines short and thin, diameter of alveoli small, and alveoli not 
so deep in females (Pocock, 1941; for Indian animals).^-** 

^^Sex related differences in the skull may be significant. In the absence of adequate 
and correctly dated material, erroneous taxonomic conclusions have sometimes been drawn. 


Fig. 93. Skull of leopard, Panthera pardus ciscaitcasica Sat. No. S 51945, 
Zoological Museum, Moscow University. Kopet-Dag, 1935. Sketch by N.N. 



165 Age-related changes in the skull of leopards are the same as those in 

tigers. In young animals the skull has a relatively short and broadly expanded 
cranium, small facial part, close-set zygomatic arches, and rather smooth 
interorbital and particularly postorbital constrictions. Crests are either absent 
or slightly developed. The skull lengthens with age. 

Age-related changes in the skull of leopards differ markedly from those 
in tigers in that their range is smaller, i.e., the raoфhological contrast between 
the early and last stages of postnatal development is less. This is explained 
by the slightly more "catlike" structure of the leopard skull and the 
elimination of the later stages of refinement seen in the tiger. Leopards 
exhibit the beginning of tliis process, wliich progressively develops in various 
species of the family and ultimately leads to a decrease in contrast between 
the early and late age-related moфhological stages. 

Geographic changes in skull structure are little known but evidently 
involve changes in overall measurements. 

Data on size of leopards inhabiting the Soviet Union^*'' are very few 
and do not cover the entire range of variation in these features. Body length 
(male and female) varies from 120 to 171 cm, tail length 75 to 102 cm, length 
of hind foot 24 to 26 cm (four animals), length of ear about 7.5 to 8.0 cm, 
and height at shoulders 50 to 78 cm. Adult animals weigh 32 to 60 kg.^^ 

Weight may possibly reach 75 kg (Stroganov, 1962). References to 117 
and even 130 kg (Baikov, 1927 and 1929), often quoted in literature (Ognev, 
1935), are positively erroneous (see below). This is underscored by the fact 
that they are said to pertain to the rather small Far East form (specimen 
from northeastern China), although representing a record for that country, 
yet fall within the range of variation of linear measurements of the species 
(body length 145 and 151 cm).^ A "very obese" male killed near Sud- 
zukhin preser/e on August 12, 1945, although measuring 128 cm in length, 
weighed only 44 kg (G.F. Bromlei). The known maximum for Amur leopard 
within the Soviet Union is 48 kg. Two Amur leopards, males of "average 
dimensions" and "moderate obesity," killed in February and March, 1927 

*4ccording to Dinnik, 1914; Satunin, 1915; Bil'kevich, 1924; Baikov, 1927; Ognev, 
1935; Stroganov, 1962; material furnished by G.F. Bromlei, N.K. Vereshchagin, A.G. 
Pankrat'ev, and A.A. Sludskii; and collections of the Zoological Museum, Moscow University 
and the Zoological Museum, Academy of Sciences. 

"^^These measurements, taken "in the flesh", cover all the published data for Russian 
animals and those obtained from skins and mounted specimens except one, i.e., 183 cm for 
a Caucasian leopard (skin from Lenkoran; Satunin, 1915). 

^^Data furnished by N.A. Baikov on the size and weight of leopard from northeastern 
China are extremely contradictory. For example, apart from the data given above, a weight 
of up to 160 kg ("iO poods"') has been quoted; 96 kg ("6 poods"; 1915) is more common. 
The length of these animals generally reaches 155 cm and the tail length not more than 
86 cm, but a figure of 95 cm has also been adduced for the same region (1927). 


near Vladivostok, weighed 32.2 and 32.8 kg (Ognev, 1935). The weight 
of a large female evidently from northeastern China was 42.5 kg; two other 
animals (female and male) weighed 25 and 29 kg (Chou, 1958). A leopard 
from Kopet-Dag weighed 60 kg; the forms there are evidently very large.^^ 

1Ъе following data provide some idea of how Russian leopards compare 
with Indian ones in size (P. p. fusca — whole of India except Kashmir, 
Nepal, Sikkim, Baluchistan, and Sri Lanka). Body length of four large 
males from India 137, 130, 128, and 128 cm, and tail length 84, 84, 76, 
and 92 cm (Pocock, 1941). Average measurements of 11 males from Sri 
Lanka: body length 126.5 cm, tail length 85.9 cm, height at shoulders 
64.3 cm, and weight about 56.34 kg (124.1 English pounds). The largest 
male from Sri Lanka (an exceptionally large animal, largest of all animals 
known) and a body length of 142 cm, tail length 96.5 cm, and weighed 
about 77.2 kg. Average measurements of seven females from Sri Lanka: 
body length 104.7 cm, tail length 77.5 cm, and weight 29 kg. The largest 
female had a body length of 114 cm, tail length 84 cm (total length 198 
cm), and weighed 44 kg (Phillips, 1935). Maximum skull length of five 
adult males from India 232 to 250 mm and condylobasal length 207 to 221 
mm; corresponding measurements in six females 177 to 201 mm and 159 to 
185 mm (Pocock, 1941). 

Greatest skull length in males 193 to 256 mm and in females 180 to 
218 mm; condylobasal length in males 186 to 223 mm and in females 170 
to 188 mm; zygomatic width in males 123 to 172 mm and in females 116 
to 135 mm; muzzle width above canines in males 53 to 65 mm and in females 
47 to 53 mm; length of upper tooth row in males 65 to 79 mm and in females 
60 to 68 mm.9o (V.H.) 

166 Systematic Position 

While lion and tiger are very closely related species, the leopard is notably 
distinct from them. From the viewpoint of systematics, the leopard is closely 
related to jaguar (P. onca). This view is supported by the coat pattern and 
craniological features. Of these two species, leopard represents a more 
specialized form (Haltenorth, 1937), i.e., a more progressive one. However, 
it is possible that leopard is closer to tiger than is jaguar, which thus 
completes the specialization series in the genus (lion — tiger — leopard — 
jaguar; Tsarapkin, 1939; see also characteristics of the genus and "Systematic 

^^''A large male up to 7 poods" (112 kg) from this region (Bil'kevich, 1924) is an 
outright misstatement. 

^^According to the data available in literature, material of the Zoological Museum, 
Moscow Univei^ity and the Zoological Museum, Academy of Sciences (N.K. Vereshchagin), 
and data furnished by A.G. Pankrat'ev (Vladivostok) and N.I. Ishadov (Ashkhabad). 


Position" of tiger). In any case the skull of leopard, compared to that of 
tiger and lion, does not exhibit many "primitive predatory" features; iastead 
it reflects a significant step toward the evolution of a distinct cat type of 
skull. (V.H.) 

Geographic Distribution 

Leopards are found in forests, parts of forest steppes (savannas), and 
montane regions of Africa and southern Asia, the Near East and the southern 
half of eastern Asia. 

Geographic Range in the Soviet Union 

The range in the Soviet Union represents the northern rim of distribution 
of the species and consists of three isolated sections — Caucasian, Middle 
Asian, and Far Eastern, which are connected to the south outside USSR 
boundaries. The Middle Asian section, in turn, is broken into a few individual 
pockets (Fig. 94). 

In the Caucasus the range (reconstructed) covers essentially the whole 
of the country except steppe areas (Fig. 95). Leopards used to occur in 
mountains and foothills covered by forests and also in desert hills devoid of 
forests. In the west the range commenced at Tuapse, possible even slightly 
more westward, but probably did not reach Novorossiisk. Intersecting at this 
meridian the western offshoots of the Great Caucasus, the range boundary 
encompassed the northern foothills and ran east and southeast up to the 
extremity of the mountain range — toward Buinaksk and Makhachkala. In 
the foothills leopards were confined to forests and at places where forests 
merged into the plain or descended to very low levels. Thus, in the western 
Caucasus leopards were known from Samursk station, Elisavetpol'sk village 
and certain other sites to the south and southwest of Maikop; in the central 
Caucasus in the region of Ordzhonikidze (Vladicaucasus); in the east, at 
Evdokimov on the Argun, near Makhachkala town itself (Agach-aul), and 
at Karabudakhkent and Gubden, southeast of Makhachkala.^^ 

^^Leopard skins sold in Kizlyar in the eighteenth century (Pallas, 1811) were most 
probably brought from the hills but quite possible leopards also intruded from the foothill 
forests into Terek valley forests, which were formerly far more extensive than at present, 
closely associated with the foothills and hill forests, and abounded in ungulates. In general, 
however, there is no information relating to the distant past (up to the nineteenth century) 
for the Caucasus; yet indubitably leopards were present everywhere in the northern Caucasus 
wherever there were forests. It is highly probable that in tiie remote past (before the appearance 
of Russians), the animal was generally far more extensively distributed in the Caucasus than 
has been recorded. Partly, as at present, these occurrences were in the form of intrusions 
along the periphery or into the plains of the eastern Trans-Caucasus, but it is impossible 
to distinguish such past instances from the fairly normal permanent habitation. 

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Along the Great Caucasus leopards occurred up to the upper forest 
boundary and even higher. On the southern slope they descended to the 
foothills and even to the plains as in the north. In the eastern part of the 
mountain range the animals were known in Alazan valley at Lagodekh, 
Zakatal, and Nukhu, and recorded in the west at Adler, Sochi, Tuapse, and 
Bzybi. They therefore descended in this region to the sea. 

In the Trans-Caucasus the range of leopards covered all of the montane 
region of Little Caucasus, including in the west the Trans-Caucasian expanse 
between Rioni and Kur', and the Pontic range in the southernmost part 
of the coast (Satunin, 1915).^2 There is no direct evidence about the 

Fig. 95. Range of leopard, Panthera pardus ciscaucasica Sat. in the Caucasus (scale in km). 

1 — probable boundaiy of reconstructed range in the historic period; 2 — region of fairly 
regular, at places very rare, occurrence of leopards by 1950; 3 — intrusions of leopards 
into the steppe; 4 — some sites where a few wandering animals were caught from the 
1930 's to the 1950's outside the territories shown by 2; 5 — possible occurrence at Terek 
in the eighteenth century and before. V.G. Heptner. 

^■^Pontic range refers here to the northernmost extremity of this mountain range to the 
south of Batumi, then a part of Russia. 


168 occurrence of leopards in the marshy lowlands and the forest expanses of 
Kolkhidsk lowland, but there is no doubt that they prowled there, though 
rarely, because of scarce game. The distribution also covers the Talysh and 
Lenkoran lowlands. 

In the steppe part of the eastern Trans-Caucasus leopards were evidently 
absent. However, wandering animals were seen even in fo restless steppes 
and semidesert plains far away from the mountains. A case of transgression 
into Sal'yan was recorded at the end of the nineteenth century (Radde, 1899) 
and one in 1946 into the densely settled part of the Apsheron Peninsula 
(Bil'gya, 25 km east of Baku; Dzhafarov, 1946); some transgressions into 
the "central Mugana reeds" have also been recorded (Vereshchagin, 1959). 

By the 1950's to the 1960's the range of leopards in the Caucasus had 
shrunk greatly, the population of the animal became negligible, and actually 
on the brink of total extinction. This was due to the ruthless killing of the 
animal, which had never been particularly abundant. The rapid and steady 
disappearance of leopards, especially in the Great Caucasus, has mainly 
occurred over the last 80 to 100 years, particularly in recent decades, since 
even at the end of the nineteenth century it was still sighted throughout 
most of the Caucasus. Not even the establishment of the Caucasian reserve 
in the 1920's could restore the dwindling population of this animal in the 
western Caucasus. 

From the 1930's to the 1950's leopards, though extremely rare, were 
still sighted in the forest regions of the westem (Kuban) Caucasus, mainly 
in the Caucasian preserve and adjoining areas. In the 1930's in this region 
there were two pockets of leopard habitation — one in the upper reaches 
of the Kuban, Kishi, and Beloe (Atamazha, Tybga, Dzhemamk, Chugush, 
and Assara ranges), and the second in the upper reaches of the Sochi, Khosty, 
Golovinka, Bzyby, and Shakha, which flow into the Black Sea. It is possible 
that a few leopards inhabited the montane regions of the eastern Caucasus 
above Zakatal and Lagodekh and possibly the heavily forested sections of 

169 Dagestan associated with them. In the rest of the Great Caucasus, on both 
northern and southem slopes, over the last decade only a few stray intnisions 
(once in several years or decades) have been reported, sometimes at places 
in the western, central, and eastern parts of the range, far from the places 
specifically mentioned above (Fig. 95)*. ^^ 

*Misnumbered "'96'' in Russian text — Sci. Ed. 

^^Occurrences at Zelenchuk and Arkhyz are known. Distant intrusions include Makhach- 
kala and the Apsheron Peninsula (see above). One leopard was killed at Lars on the Georgian 
Military Highway in 19()0 (Beme, 1941) and another roughly at the same place in the 1940's. 
In February, 1949 a pair of leopards was seen at Galaskha in the Groznyi region. Other 
instances of distant intrusions are known, for example, towards Tskhinvala (soutiiem Ossetia, 
1952) (see also Vereshchagin, 1959). 


In the Trans-Caucasus (not including the Great Caucasus range), in 
the 1950's and early 1960's leopards still inhabited the Talysh, Karabakh, 
and Zangezur ranges in the north to the Kel'badzhar region and Murovdag, 
and westward along the mountains to the north of the Araks valley through 
Nakhichevan Autonomous Soviet Socialist Republic almost to Yerevan (to 
the Saraibulag range and the Veda region). They also occurred in Kubatlinsk 
and Zangelansk regions of Azerbaidzhan and possibly from time to time 
in the northern forest regions of Armenia (north and northwest of Lake 
Sevan). This section of leopard habitat, at least in its southern portion, was 
associated with the geographic range in Iran and intmsions southward 
occurred from there. The probability of such intrusions has diminished in 
recent decades. In the mid- 1960 's leopards were already absent in Armenia, 
except probably for some individual animals in the Zangezur range. On 
the Talysh, however, leopards were resident throughout the 1960's and 
evidently not mere intruders. Almost every year one or two leopards are 
killed there to the west of Astar and Lenkoran (road to Lerik). Two animals 
were shot in October, 1969 west of Astar (V.S. Lobachev). 

On the whole, by the middle and end of the 1960's leopards had already 
practically disappeared or were passing through their last days in the Trans- 
Caucasus and the Little Caucasus. They still occur, though very rarely, at 
places where there are several tributaries from Iran, i.e., on the Zangezur 
(southeastern Armenia and southwestern Azerbaidzhan) and persist on the 
Talysh. Some animals may have also intruded from these areas into other 
parts of the Trans-Caucasus. Leopards do not enter the Soviet Union from 

In the Great Caucasus some leopards are evidently still met with along 
the southern slope of the eastern section (Azerbaidzhan and eastern Georgia) 
and some strays probably live in the Kuban region of the Caucasus and 
on the southern slope of the western half of this range. Leopards are absent 
in the rest of the Great Caucasus and in certain areas have not been seen 
for quite some time.'-''^ 

The Middle Asian habitat of leopards (Fig. 96) in the Soviet Union is 
associated with the distribution of the animal in the Near East, in Iran and 
Afghanistan, and represents the northern extremity of its range. It is divided 
into several sections. In the west, in Turkmenia, a large section covers all 
of the Kopet-Dag from its western and northwestern (Kyuren'-Dag) rim 
to the extreme east, i.e., roughly to Artyk railway station (about 59°30' E. 

^'^The range in the Caucasus, in addition to works cited in the text, is based on Dinnik, 
1914; Satunin, 1915; Heptner and Formozov, 1941; Nasimovich, 1941; Vereshchagin, 1947 
and 1949; Burchak-Abramovich and Dzhafarov, 1949; Dal', 1954; Kotov and Ryabov, 1963; 
Alekperov, 1966; and other sources; as well as original data compiled by V.G. Heptner. 

See section "Biology"" for more details about the Caucasus and the reduction of the 
range there. 


long.), where the mountain extends beyond the Soviet Union. An isolated 
very small section is formed by the Little Balkhan upland (774 m above 
sea level) and a slightly larger one by the Great Balkhan (1,880 m above 
sea level). Both of these are isolated from each other and from the Kyuren'- 
Dag section by desert plains — the so-called Balkhan passes ("corridors"), 
each 25 km wide. Obviously, the leopard population in these sections is 
likewise isolated. Yet, in spite of the Turkmenian leopard's affinity for 
mountains (it does not enter the plains for more than a few kilometers), 
one could nevertheless assume that formerly, when leopards (like mountain 
sheep) were widespread in Kyuren'-Dag and their population high there, 
some exchange of animals must have occurred between Kopet'-Dag and 
the Balkhans (Fig. 96). 
170 Farther to the east there is a separate section (within the Soviet Union) 

of leopard habitat to the west of Badkhyz in the Gyaz'-Gyadyk upland (up to 
1,000 m) extending from Iran. There the animal reaches in the north, to 
Pul'-i-Katum on the Tedzhen and is found in the middle of Badkhyz in 
the Lake-Er-oilan-daz depression (ur. Namak-saar) and in the Kyzyl-Dzhar 
gorge. Farther east, again after some intermption, leopards occur in the 
mountains east of the Kushka River, between Kushka and Kashan and the 
upper Murgab. Toward the north there they are not found beyond the mouth 
of the Kushka on the Murgab and probably do not even intmde in that region. 

Information about the habitat of leopards along the Tedzhen and Murgab 
outside the sites discussed above is erroneous and mainly due to mistaking 
cheetah for it. North of the Great Balkhan leopards are not encountered 
and occasional information about their occurrence there also pertains to 
cheetah. In the Kopet-Dag leopards occurred (or still did even in the 1940's) 
right up to the foothills of the range and sometimes even emerge onto the 
premontane plain. Thus, intruding from Iran, they stayed from time to time 
in the premontane plain slightly east of Artyk, even at Mean and Chaach 
(south of Tedzhen city) and moved toward the Caspian Sea in the Atrek 
basin. Similarly, in southern Turkmenia leopards from Gyaz'-Gyadyk went 
east into the Er-oilan-duz depression (data for Turkmenia from V.G. 

A separate section of the Middle Asian range of leopards is situated on 
the right bank of the upper Amu-Darya and Pyandzh. There leopards live 
in low (up to 1,100 to 1,400 m above sea level) mountains in the west up 
to Babatag (between Surkhan-Darya and Kafirnigan), in the north to the 
Gissar valley (upper reaches of the Kafirnigan), and in the east to the foothills 
of the Darvazsk range and the Parkhar region on the Pyandzh, where their 
range adjoins that of the ounce [snow leopard]. Within the boundaries 
described above, leopards have been sighted in Bahatag, Aktau, Oktau, 
Karatau, Terglitau, the Bal'dzhuan mountains, in the regions of Koktash, 


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Shaartuz, Dangarin, Mikoyanabad, and Parkhar (Flerov, 1935; Chernyshev, 
1950; Ishunin, 1961). 

All information about the residence and catch of leopards in Semirech'e 
(Shnitnikov, 1936; and others) is erroneous and refers to ounce. Even random 
intrusions of leopards into this part of Middle Asia should be regarded as 

171 Fig. 97. Range of leopard, Pantherapardus orientalis Schleg. in the Far East (scale 

in km). 

A — northern boundary of fairly permanent habitation in the 1950's to the early 
1960's (data of G.F. Bromlei; N.V. Rakov, 1965); B— boundary of region of 
maximum permanent habitation in 1950 (G.F. Bromlei); С — intrusions in the 
middle of the last century (1 — Tynna River, and 2 — Bureinskii mountains); D — 
some intrusions in the 1910's to the early 1920's (3— upper reaches of Iman, and 
4— Sitsa); E— some intrusions from 1939 to 1962 (5-30 km south of Vyazem, 
6 — left Pokkhorenok, and 7— Bikin midreaches). Section of the course of the 
Amur between "a" and "b" represents region of leopard intrusions in the middle 
of the last century (Maak, 1859). For intrusions of leopards into Trans-Baikal, 
see Fig. 94. V.G. Heptner. 


In Middle Asia the reconstructed range of leopards described above, 
unlike the situation obtaining in the Caucasus, mostly still corresponds to 
171 the range at the end of the 1960's. Leopards, however, are absent in the 
northwestern part of the Kopet-Dag (in Kyuren'-Dag) and became extinct, 
or almost so, in the Great Balkhan by the early 1950's. The population 
throughout the range also underwent radical changes (see below). 

The Far East range of leopards (Fig. 97) within the Soviet Union is 
contiguous with its range in northeastern China and the Korean Peninsula. 
The predator has been found in the Ussuri territory and in parts of the Pri- 
Amur and Trans-Baikal. 

In the middle of the present century (1940's to 1960's) the region of 
fairly permanent habitation, including regular intmsions (see below) was 
limited to the far south of Ussuri region. The northem boundary of the range 
there commences on the coast of the Sea of Japan and Dzhigit Bay at 44° 
N. lat. and runs south at a distance of 15 to 30 km from the coast to the 
latitude of Valentine Bay (43°10' N. lat.). There it turns steeply westward, 
north of the Suchan basin, then north to encompass the source of the Ussuri 
(Daubikhe River) and its right bank tributary, the Ulakhe, part of the basin 
of the right tributarj' of the Ulakhe Noto, and the Shetukhe, the right tributary 
in the upper reaches of the Ussuri (G.F. Bromlei, Rakov, 1965). Here the 
boundary turns westward toward the bank of [Lake] Khanka at the level 
of the state boundary (Figs. 97 and 98). 

South and west of the above line leopards occur everywhere, at least, 
172 they did in the past, in the Khanka lowland (south of the lake). The asso- 
ciation of Ussuri leopards with mountains is fairly definite but evidently 
not as rigid as in the case of Turkmenian leopards. In the area described 
leopards are rare, especially in the more northern parts; they are confined 
more to places where wild sika deer live or where deer husbandry is 
practiced. Thus in the 1950 's leopards were observed at Razdol'nii (50 km 
north of Vladivostok), Gamov Cape, Kita Bay, "Kedrovaya Pad' " preserve, 
Tuman Bay, Glazkov, along the rivers Steklyanukha, Maikhe, and Lefa, 
and elsewhere (G.F. Bromlei; for more details, see also Rakov, 1965). 

The unique feature of leopard habitat in this part of the Soviet Union 
is that this area is one of occurrence of solitary wandering animals, groups 
in pairs, or a mother with cubs. Definite data on breeding of leopards 
throughout this region are not available and it is possible that the animal 
does not reproduce in this part of the Soviet Union except perhaps in 
"Kedrovaya Pad' " preserve (southwest of Vladivostok). In general, it 
consi-sts of animals originating from the western"' part of China. This does 
not contradict the fact that here they are a settled species and relatively 

*S/c: should read "eastern" — Sci. Ed. 


Anyui R. 

ППШПав^ 4 


Fig. 98. Present distribution of leopard, Panthera pardus orientalis Schleg., in 

Ussuri territory. 

1 — boundary of region of permanent residence by 1971; 2 — best sites of 

habitation; 3 — region of very rare and temporary intrusions beyond zone of 

permanent occurrence; 4 — directions of individual intrusions from China during 

1930 to 1970. (S.P. Kucherenko.) 

common in places. Such a position is generally quite normal for the periphery 
of the range of distribution. This explains, probably, some ambiguities in 
establishing the boundaries of the leopard's range in the southern Ussuri 
region. These, however, pertain to frequencies of sightings. A comparison 
of range data for the past and present brings us to the conclusion that the 
region of permanent habitation of leopards in the Ussuri territory has not 
altered significantly over the last hundred years. 

With such an insignificant region of more or less regular habitat leopards 

cover a large area with long intrusions which, at one time, contributed to 

173 an incorrect depiction of the range of the species in the Soviet Far East and 

in Trans-Baikal. This is mostly explained by the fact that older zoologists 


usually relied on inconsistent information from local people and were partly 
influenced by the erroneous statements of Pallas (1811) about ounce between 
the Uda and Amur.^'^ Thus it was even thought that leopards inhabited the 
Amur region to the same extent as tigers, i.e., the entire course of the Amur, 
the shores of the Okhotsk and Tatarskii seas, and Sakhalin Island (Schrenk, 

At present there is justification to consider all the areas where leopards 
have been found outside the region described above as regions representing 
deep and rare intrusions. Furthermore, all these areas represent intrusions 
not from the south of the Ussuri territory but from northeastern China. Only 
the intrusion into the Sitsa River region may be an exception. The following 
are the northernmost known points of leopard occurrences: southeastern 
Trans-Baikal (very rare); Kuchugai on the lower Gazimur, a tributary of 
Argun in eastern Trans-Baikal (Cherkasov, 1887; Baikov, 1927); Nerchinsk- 
Zavodsk region of Chitinsk district (1952; A.A. Sludskii); Bureinskiy 
mountains (at the Amur; Radde, 1862); Tyrma River, a tributary of 
the Bureya (evidently in the 1фрег reaches in these same m.ountains; 
Middendorff, 1867); and along the Amur between the mouths of the Sungari 
and Gorin.^^ All these represent very distant and, evidently, rare intrusioas 
in the past century, in part in the middle or the first of it. 

In the 1910's and 1920's intrasions of individual animals were observed 
in Sitsa in the Terneya region (coast of the Sea of Japan in the proximity 
of 45° N. lat.) and in the upper reaches of the Iman (around 46° N. lat.; 
Arsen'ev, 1926; Baikov, 1927). In the middle of the present century 
intrusions are known in the middle course of the Bikin, the Khor basin 
(the Lxft Podkhorenok), on one of the right tributaries of the lower reaches 
of the Ussuri (30 km south of Vyazem.), to the mouth of the Bir and on 
its middle course (Rakov, 1965), and others [sic]. In the past, intrusions 
were evidently more frequent and extended farther, but even at present 
leopards move quite far, even in Trans-Baikal (additional data on intrusions 
and distribution in the range are given under "Biology"). 

Old information about the habitat of leopards in Sakhalin (Schrenk, 1858; 
Nikol'skii, 1889) is undoubtedly erroneous. Similarly, assumptions about 
leopard habitat in the eleventh and twelfth centuries at the mouth of the 
Don and in the southern Russian steppes or forest-steppes, based on an 
inteфretation of the "fierce animal" of old Russia as leopard (Pidoplichko, 

^•''Older researchere (Maak, Schrenk, PKheval'skii, and Nikol'skii), following Pallas, 
usually applied the name Felis irbis to the Far East leopard. This has led to confusion even 
among some present-day zoologists (Zhitkov, 1934) and is a purely nomenclatural enor. 
Ounce are absent in the Far East and were never present there; all data pertain to the leopard. 

^^In the expanse "from the mouth of the Sungari ending slightly above the mouth of 
the Gorin, the Tuzemtsy [local people] were aware, apart from tiger, of still another big 
species of the cat genus"' (Maak, 1859). 


1951; Mavrodin, 1964; and others), are unfounded (see section on tiger 
and also Heptner, 1969). The discovery of parts of the lower jaw of a leopard, 
used as an amulet, in the ruins of one of the Greek Black Sea colonies 
cannot serve as adequate proof (Vinokur and Khotyun, 1967). This should 
merely be considered an import from Asia Minor, with which Black Sea 
Greeks conducted an active trade. 

On the whole the range of leopards in the Soviet Union is in the process 
of undergoing reduction, at places very rapidly. The population is partly 
supplemented by animals migrating from adjacent territories, where they 
are more common and suffer less persecution (Iran, Afghanistan, and China). 
If measures are not implemented soon for the conservation of leopard, this 
species will totally vanish from the Soviet Union in the near future. 

Geographic Range outside the Soviet Union 

The range outside the Soviet Union includes northeastern China (Manchuria), 
the Korean Peninsula, eastem China (evidently not Kansu), the eastern border 
of Tibet,^^ Indochina, Malacca, Java, Kangean Islands,^^ Burma, India, 
174 the Himalayas, Pakistan, Afghanistan, Iran, Asia Minor, the Arabian 
Peninsula, Sinai, Egypt, Morocco, Algiers, Tunisia, and the whole of Africa 
south of the Sahara (Fig. 99). 

At present the range everywhere is drastically reduced. 

In the Quaternary period leopards lived in France, Italy, England, 
Germany, Belgium, Spain, Portugal, Switzerland, Yugoslavia, Hungary, and 
Rumania. Some discoveries point to the occurrence of this species in Europe 
up to and including the Neolithic period (Pidoplichko, 1951). Within the 
USSR leopard remains have been found in the JVIiddle Quaternary formations 
of the Tashik-tash caves in the Baisunsk mountains, which represent the 
very fringe of the present-day range. In strata of the fifth to the second 
century B.C. a lower jaw was found at Olvia (Nikolaevsk district). This 
was evidently the remains of an imported animal. (V.H.) 

Geographic Variation 

Geographic variation in leopards is well-marked, wliich is natural considering 

^^In general, usual references to "Tibet" are erroneous. Leopards are evidently not 
seen in the uplands and in the ranges surrounding them from the north. Residence in eastem 
Tibet (Kani and others) has been established with certainty. 

^^In Sumatra leopards are absent and all data about their residence there should be 
considered erroneous. At the same time the break in the range at Malacca-Java appears very 
strange and probably leopards were present in Sumatra in oiden times. The existence there 
of a very small number of animals has been assumed even by recent authors (Chasen, 1940). 

References to the occuirence of leopards in Japan (DaP, 1954) are unsubstantiated. 



Fig. 99. Reconstructed range of leopard, Panthera pardus Satunin (scale in km). 

In the northeast the boundary also includes region of intrusions. Information about 

Sumatra appears dubious. V.G. Heptner. 

the vastness of their range and the diversity of natural conditions within 
it. Nevertheless this variability is not as shaф as might be expected under 
these conditions, although extreme forms differ rather notably. The main 
background color of the coat may be very pale — pale yellow or even 
grayish-white — on one side of the range, and bright — red or reddish- 
chestnut on the other. A transition between these extremes evidently-does 
not occur 8Ьаф1у anywhere, although in some parts of the range a subspecies 
or a different category lives in relative proximity. 

In other features variability is reflected in intensity of the black tone 
of the spots, their number, size, density of disposition, relative number of 
rosettes and their position on the body, the color of the center of these spots, 
etc. For some forms a dual-color type has been noted. In some populations 


of the humid regions of southern Asia the percentage of melanistic forms 
("black panthers'') is high (Sikkim and Nepal — P.p.pernigra and Java — 
P. p. melas), while in other predominantly arid zones melanism is totally 
absent or an extremely rare phenomenon. 

On the whole the geographic variability of leopards distinctly conforms 
to Gloger's rule. This pertains not only to the general shade of color, but 
also to the total area of black patterning and number of melanistic forms. 
Within the USSR and adjacent sectors of Asia, Bergmann's rale is not 
manifested so distinctly, but in the range as a whole much smaller forms 
are associated with warmer and more arid regions. At the same time, for 
example in East Africa, these animals are very large, judging from the skull 
size. Such big animals have not been reported in the Soviet Union to date 
(Best et al., 1962). 

Geographic variation in leopards has not been studied adequately. At 
present a large number of subspecies have been described and the number 
of distinct and even accepted forms far exceeds those actually existing. 
Almost all the subspecies described are based on color characteristics; 
however, as pointed out before, coloration exhibits extreme individual 
variability. Individuals exhibiting extreme color variability from a given 
region have sometimes been placed in separate subspecies (see later). 
Geographic variation for the entire species has not been revised in accordance 
with our present level of knowledge. 

At the same time contravening modem concepts (also in nomenclature), 
the practice continues of maintaining as distinct old forms and even separat- 
ing new ones on the basis of casual data and even stray photographs and 
skin descriptions. Disagreements and changes in the concepts of geographic 
variation of the species are coasequently great indeed. Hence 15 subspecies 
were recognized from Africa in the 1930's (over 25 names; G. Allen, 1939); 
of these, many pertain to the eastern part of the mainland and three to south 
Africa (Roberts, 1951). Later (Ellerman, Morrison-Scott and Hayman, 1953), 
all South African animals were placed in one nominal form (Egypt), i.e., 
the African expanse from north to south is now regarded as populated by 
one single subspecies. This positive critical approach contradicts the tendency 
of extreme subdivision. A large number of leopard subspecies are recognized 
from Asia (12; Ellerman and Morrison-Scott, 1951 and 1966). 

In the Soviet Union various zoologists have generally designated five 
subspecies. In reality only two exist. 

1. Amur leopard. P.p. orientalis Schlegel, 1857 (syn. villosa, japonensis, 
fantanieri \sic\ fontanieri], chinensis, grayi) 

Size not large. 

Coat fairly soft, with long (on the back 30 to 50 mm and on the abdomen 
70 mm) and dense hair. Main general color type bright and lustrous. Winter 


coat varies from fairly light yellow to dense yellowish-red with a golden 
tinge or rusty-reddish-yellow. Color on flanks and outer sides of legs lighter. 
Spots pure black color; light -colored centers of circle of spots ("rosettes") 
somewhat darker than main background color of skin. Spots numerous, i.e., 
spottiness prominent (Fig. 100). Summer pelage shorter and brighter with 
more vivid coloration pattern. 

Skull small, with narrow interorbital region (on average, width about 
20% of condylobasal length); postorbital constriction distinct, short, and 
in the form of an isthmus; nasal pointed at posterior ends and zygomatic 
arches relatively massive. 

Measurements of male (six): body length 107 to 136 cm (M 125); tail 
length 82 to 90 cm (M 86); length of hind foot 24 to 27 cm (M 25); and 
height at shoulders 64 to 78 cm (M 72). 
176 Maximum length of skull 204 to 232 mm (M 213); condylobasal length 

186 to 200 mm (M 189); zygomatic width 129 to 144 mm (M 138); 
interorbital width 34.3 to 39.9 mm (M 37.8); postorbital width 36.8 to 45.0 
mm (M 40.2); and length of upper tooth row 67.8 to 68.7 mm (M 68.0) 
(Stroganov, 1962). 

Weight of males of moderate size 32 kg (Ognev, 1935) and of large 
ones 48 kg. This weight may even reach 60 to 75 kg.^^ 

This subspecies is encountered in the Amur and Ussuri regions. 

Outside the Soviet Union Amur leopards are found in northeastern 
China, probably in the south to Peking, and the Korean Peninsula. 

Amur leopards are clearly distinguishable from leopards inhabiting 
Turkestan and the Caucasus. 

The differences between Amur and Chinese leopards are complex and 
not fully understood. Thus some false notions have arisen. As mentioned 
above, the range of color variability in Ussuri leopards is quite broad and 
differences in color intensity and brightness of extreme forms fairly sharp. 
Differences in vividness of coloration between the winter and summer coats 
are also distinct. This sometimes creates the impression of the existence in 
the Soviet Far East of two forms of leopard (subspecies), differing in color. 

^^In northeastern China (Manchuria) out of 18 animals killed and 18 skins the 
following are the measurements of the two largest (Baikov, 1927 and 1929): body length 
145 and 151 cm, tail length 95 and 82 cm, and maximum length of skull 221 and 255 mm. 
The weight of these two animals has been shown as 117 and 130 kg which, of course, appears 
erroneous (see "Description"' above); this is the weight of adult Amur tigresses. (The author 
perhaps meant pounds and wrote kilograms by mistake — ? On converting Russian pounds, 
the weight would be 46.8 and 52 kg and English pounds 53.0 and 58.9 kg, which in general 
accords with other data.) The following are the dimensions of three adult males from 
northeastern China: body length 106, 110, and 106 cm; tail length 76, 70, and 70 cm; length 
of hind foot 23, 22, and 24.3 cm; height of ears in all three animals 7.5 cm; and weight 
25, 42, and 29 kg respectively (Shaw, 1958). 





TO ._ 




uT с 





The presence of bright-colored specimens has provided some authors 
(Satunin, 1914; Bobrinskii, 1944; Novikov, 1956) a basis for suggesting 
the occurrence or even regular occurrence of Chinese leopards (P. p. 
ch'mensis) in the southern Ussuri region. 

These same two major color types of leopard have also been established 
for northeastern China (Manchuria) (Baikov, 1927, 1927a, and 1929), where 
they have been designated distinct subspecies ("Manchurian" orientalis 
and "northern СЫш" fotitanieri). There is no doubt that these represent 
only color variants in China, but the boundaries of distribution of the 
subspecies orientalis in the south are not well defined. Whether there is 
(or was) in fact an independent form chinensis (fontanieri), described from 
around Peking, or animals from there belong to orientalis, or are identical 
with one of the southern forms, or represent a transition from the northern 
forms to the more southern, is not clear at the present time. Evidently, 
however, the same form of leopard exists within the Soviet Union as occurs 
in northeastern China and the Korean Peninsula. 

2. Caucasian leopard. P.p. ciscaucasica Satunin, 1914 (syn. saxicolor 
transcaucasica; the name tulliana is commonly used in Soviet literature 
for this form). 

General size, on average, somewhat larger than that of Amur leopard. 

Color of winter coat very light and pale. Main background color pale, 
grayish-ocherous, sometimes light gray with sandy or various intensities 
of reddish, but invariably relatively faint. Color more vivid on back. 
Sometimes background color even grayish-white, being similar to the main 
color shade of the ounce. Spots relatively few, usually not pure black, and 
often with a brownish tinge. Light-colored centers of rosettes usually not 
darker than main background of coat. Summer coat light, being lighter and 
less dense than that of Amur leopard. Individual color variability significant; 
two main color types distinguishable — very light and very dark. 
178 Background in light-colored type ordinarily very light grayish-ocherous 

with a faint reddish tinge; on the back, especially in the anterior part, 
slightly darker. Sometimes this type is extremely light, i.e., whitish with 
a light yellow tinge. Most of the spots are compact and relatively small 
(about 2.0 cm in diameter). Rosettes consist of three to five spots. On 
the thighs these spots quite often form two bracket-shaped rosettes turned 
toward each other. Three or four almost complete black rings occur at the 
end of the tail. Along the middle of the back, in the region of the sacrum, 
two rows of large (about 4.0 cm long and 2.5 cm wide) elongated spots 
are present. 

Background color in dark-colored type ordinarily very dark and reddish. 
Spots larger (compact ones about 3.0 cm in diameter) and sparser; most of 
those on the sacmm about 8.0 cm and 4.0 cm in size. A significant number 


of the rosettes form complete rings. Transverse markings present throughout 
most of the tail (Rerov, 1935, with modifications)/'^'^ 

Skull evidently somewhat larger than that of Amur leopards, interorbital 
region relatively broader (about 22% of condylobasal length), and region 
of postorbital constriction elongated and lateral sides almost parallel (short 
post-orbital "isthmus'' absent). Nasals relatively moderately narrowed 
posteriorly and zygomatic arches relatively less curved. ^'^^ 

Body length 126 to 171 cm (183) and tail length 94 to 116 cm.^'^^ 

Maximum length of skull in males (six) 202 to 256 mm (M 232.5) 
and in females (two) 201 to 218 mm; condylobasal length of skull in males 
(four) 185 to 223 mm (M 206.5) and in females 186 to 188 mm; zygomatic 
width in males (seven) 133 to 172 mm (M 156.5) and in females 122 to 
135 mm; interorbital width in males (four) 41 to 51 mm (M 47.2) and in 
females 41 mm; width of rostral portion above canines in males (four) 50 
to 65 mm (M 58.2) and in females 50 to 53 mm; and length of upper tooth 
row in males (four) 68 to 75 mm (M 72.0) and in females 64 to 67 mm. 

Weight may reach 60 kg (Bil'kevich, 1924).^'^^ 

This subspecies is found in the Great Caucasus, Trans-Caucasus, 
southern Turkmenia, and southern Tadzhikistan. 

Outside the Soviet Union Caucasian leopards are found in Iran, 
Afghanistan, and western Pakistan (Baluchistan). 

Differences between leopards inhabiting Turkestan and the Caucasus 

^•^It has been stated that there are almost no transitional forms between these color 
t>'pes (Flerov, 1935). Such a view is based on a small amount of data. In fact, these represent 
only extreme forms of a continuous series of variations. It is possible that one or the other 
type may predominate in different parts of the range. Thus, two color "phases" of forms 
pertaining to P. p. saxicolor are shown for northern Iran. In one, distributed in Astrabad 
and Mezanderan, the main shade is more gray than yellow and sprets more dark brown than 
black; spottiness on trunk shows distinct rosettes; lower portion of flanks, shoulders, and 
thighs with annular spots; and pattern on head small, mostly rounded spots. More to the 
east, in Kopet-Dag, Aladag, and the "Mastschid" [Meshed? — V.H.] mountains, "the color 
of leopards is more yellow than gray. Spots pure black. Annular spots do not forai rosettes 
[i.e., not broken — V.H.]. Lower part of flanks, shoulders, and rear of thighs covered with 
solid spots. Spottiness on head intense and spots irregular, sometimes angular" (Tsukovskii, 
1959). Some details of these characteristics as also references to the localization of "phases" 
give rise to doubts but on the whole the data are correct. 

^'^^The craniological features of the subspecies (according to Ognev, 1935; Stroganov, 
1962) are only tentative and need revision based on new material. 

^''^According to data available in literature (Dinnik, 1914; Satunin, 1915; Ognev, 
1935). Measurements mainly of skins and stuffed animals (see "Description"). The biggest 
leopard measured "in the flesh" had a body length of 171 cm along the curvature of the 
back, and a tail length of 102 cm (Kopet-Dag; Bil'kevich, 1924). Craniological measurements 
are from the same sources and from material of the Zoological Museum, Academy of Sciences 
and the Zoological Museum, Moscow University. 

^"^See also "Description". 


and those of the Far East are factual even though described inadequately 
(Fig. 101). However, the relations between populations from different parts 
of the above regions and those of surrounding regions are far from clear. 
The subspecies name tulliana has long been assigned to leopards of the 
Trans-Caucasus and Turkestan; this name underscores their identity with 
the form described in the middle of the last century from the western fringe 
180 of Asia Minor. Later (Satunin, 1915), the leopard of the Great Caucasus 
{P. p. ciscaucasica) was separated from the Trans-Caucasian form based 
on material from the Kuban Caucasus. Animals living east of the Caspian 
Sea were usually placed among the Trans-Caucasian forms. This scheme 
(Satunin, 1914; Ognev, 1935) prevailed until recently, even though the form 
saxicolor had been described from Astrabad by the end of the 1920's 
(Pocock, 1927), which should logically include leopards of Turkmenia, 
Tadzhikistan, and the Trans-Caucasus. This has been partly suggested by 
the author's description who thought that even the Kuban Caucasus leopards 
belonged to the form saxicolor }'^'^ 

The scheme adopted here is that leopards of Turkestan, the Trans- 
Caucasus, and the Great Caucasus are identical not only with each other, 
but also with leopards of Iran (at least northern Iran).^°-'* This single form 
should bear the less convenient name used here, which has priority over 
saxicolo?'. The latter may have importance only when the independence 
of the subspecies of Great Caucasian leopard has been demonstrated. Such 
a possibility, however, is extremely unlikely, partly because the animal in 
the Great Caucasus has been destroyed, or almost so, and is on the verge of 
extinction. It is difficult to establish the relationship of the leopard population 
of the above territories with the Asia Minor form tulliana. Evidently, 
however, it is more distinct than the form designated saxicolor (i.e., 
ciscaucasica) and the boundary of separation between these forms mns 
through Mesopotamia (Pocock, 1941).^°*^ In spite of all this, it is quite 
possible, that leopards of Asia Minor, the Caucasus, Iran, Afghanistan, and 
Middle Asia represent identical subspecies and hence should be termed 
tulliana. In the face of this possibility the recognition of four subspecies 

^'•''The fomi transcaucasica, described by Tsukovskii (1964), is not discussed here. 
It has not only been separated by upsetting prevailing procedures, but its range, according 
to the author, is not clear. Thus he exposes his ignorance not only of the distribution and 
population of leopards in the Caucasus, but also the geography of this country'. It is indisputably 
a synonym for the Middle East fomi no matter what name is assigned to it. 

105рщ. tij,(jei-5 niake a clear distinction between Far East and Middle East leopards, 
but do not differentiate Caucasian from Turkmenian leopards (Kuznetsov, 1941). 

^•^'^he fomi tulliana, formerly widely distributed in Asia Minor, is also few in number 
and its range has shrunk sharply. Its occurrence in the 1940\s and 1950's was confined to 
the extreme southwestern comer of the peninsula between the lower course of the Chediz 
and Antalya Bay (Kosswig, 1955) or only over a slightly wider area (Kummerloeve, 1967). 


179 Fig. 101. Variation in color in Turkmen leopard, Panthera pardus ciscaucasica 


1 — No. 53348, Kharasangli gorge, Kopet-Dag, April, 1948, winter coat; 2 — No. 
29657, Turkmenia, winter coat; 3— No. 438920, Sheriauk, Kopet-Dag, October, 
1941, winter coat; 4 — No. 29656, Trans -Caucasus, summer coat. Scale same. All 
skins from collection of the Zoological Museum, Moscow University. Photograph 

by V.G. Heptner. 


of leopards (saxicolor, sindica, dathei, millardi; Kullmann, 1967-1968) for 
Afghanistan alone is highly improbable. Apart from light -colored animals 
of the type ' ' saxicolor,' ' the occurrence there of the Kashmir form, millardi, 
is possible, but no other. 

Outside the Soviet Union many subspecies have been described, numbering 
over 50 names, and new ones are continually being added. The number 
of actual subspecies is evidently far less than those recognized by most 
zoologists (about 25 to 30). Recently, apart from the two forms described 
above, which are spread far beyond the Soviet Union, the following forms 
have been recognized, although significant differences of opinion exist in 
this regard (Haltenorth and Trenze, 1956, with some modifications). 

Asia: 1) P.p. minor Hemp, and Ehrenb., 1833 — Pale.stine, Syria, and 
Arabian Peninsula; 2)P.p.jarvasi Pocock, 1932 — Sinai; 3)P.p. tidUana 
Valenc, 1857 — Asia Minor; 4) P. p. sindica Pocock, 1930 — eastern 
Baluchistan and Sind; 5) P. p. millardi Pocock, 1930 — Kashmir; 6) P. p. 
pernigra Gray, 1856— Nepal and Sikkim; 7) P. p. fusca Meyer, 
1794 — whole of the Indian Peninsula except Sind, Kashmir, Nepal, and 
Sikkim; Sri Lanka, Burma, and southwestern China (from Szechwan to 
Fukien); 8) P. p. delacouri Pocock, 1930 — Indochina, Siam, and Malacca; 
9) P. p. melas G. Cuv., 1809 — Java and Kangean Islands; and 10) P. p. 
japonensis Gray, 1862 — northern (not northeastern! — V.H.) China. 
181 Africa: 11) P. p. panthera Schr., 1777 — Morocco, Algiers, Tunisia, 

Senegambia, and western Sudan; 12) P. p. pardus Linn., 1758 — Egypt, 
eastern Sudan, and to Tanganyika; 13) P. p. andnorii De Beaux, 
1923 — Eritrea and Ethiopia; 14) P. p. adiista Pocock, 1927 — high mountains 
of southern Ethiopia; 15) P. p. brockmani Pocock, 1932 — northern (formeriy 
British) Somali; 16) P.p. nannopardiis Thomas, 1904 — southern (formerly 
Italian) Somali; 11) P. p. adersi Pocock, 1932 — Zanzibar; 18) P./?. chid 
Heller, 1913 — Uganda; 19) P. p. leopardus Schreh., 1777 — forest regions 
of western Africa from former Spanish Guinea to Nigeria; 20) P. p. 
reichenovi Cabrera, 1918 — steppes of northern Cameroon and the region 
south of Lake Chad to Congo forests; 21) P. p. itiirensis Allen, 
1924 — forests of Cameroon and Congo; 22) P. p. shortridgei Pocock, 
1932 — Angola and Damaraland to Mozambique; 23) P. p. puella Pocock, 
1932 — Caocoveld, southwestern Africa; and 24) P.p. melanotica Gunther, 
1885 — Cape Colony to southern Rhodesia. 

The above forms, especially those of Africa, can be combined into 
groups based on color characteristics and associated with specific natural 
and geographic conditions. For example, deep-colored animals occur in wet 
tropical forests and light-colored ones in savannas, while even lighter-colored 


ones live in desert regions. Quite possibly these groups in fact represent 
actual subspecies. While some contemporary researchers have a tendency 
to split groups, others tend to just the opposite. Thus all the four forms 
of southern Africa (Nos. 19, 22, 23, and 24 of the above list) have been 
combined into one group and recognized as identical with the Egyptian 
(nominal) form, i.e., it is assumed that most of Africa is inhabited by a 
single form (Ellerman, Morrison-Scott and Hayman, 1953). (V.H.) 


Population. Until the end of the last century leopards were found almost 
everywhere in the mountains and foothills of the Caucasus., but absent by 
and large in plains and densely inhabited regions. Leopards were relatively 
common in the northwestern extremity of the Great Caucasus range. In 1893 
they were sighted even at Tuapse and somewhat earlier (in 1875) twice 
at Sochi (Radde, 1899). From time to time they have been met with north 
of Fisht and Oshten, not far from Samur station on the Pshekha, a tributary 
of the Beloe, around Elizavetpol'sk village, and at other places (Dinnik, 
1914). In the 1880's leopards were killed on Dudugash mountain (lower 
Kishi [river]) and occurred in 1895 around Slesamaya mountain (Shisha 
River) (Rossikov, 1890; Nasimovich, 1941). In 1896 they were caught on 
the upper Kishi and they were seen a year later on the upper Sakhrai and 
on the Pshekisha (Dinnik, 1914). North of Abago (on the upper Beloe) in 
1895 N.Ya. Dinnik saw two leopards in the course of one day, and in 1907 
in the course of two days thrice heard their roar at midday in the mountains 
between the Kishi and the Urushten. According to him formerly leopards 
were caught quite frequently above Psebai in Malyi Laba basin between 
Umpyr and Zatish'ya settlements. Thus in this region 11 leopards were 
killed from 1894 to 1896, 2 in 1904, and none in 1905 and 1906. 

At the end of the last and the beginning of the present century, in the 
Malyi Laba and UriLshten basins, leopards were encountered at many places 
but evidently confined mainly to the Great and Little Balkhan mountains 
(Nasimovich, 1941), and in the winter of 1896 to the upper reaches of the 
Andryuk (Dinnik, 1914). Around 1903 this predator was shot in the Nikits 
sanctuary in Lysaya mountains. In the Bol'shoi Laba basin all of the old 
references to the presence of leopards pertain to the region of the Beskes 
River and Markopidzh mountains. Leopards were sighted quite frequently 
on Beskes River (Nasimovich, 1941). Around 1905 a leopard was reported 
as present between the Oshtensk and Guzeripl' mountains, and some years 
later near Nagai-Koshka mountain. In 1913 and 1914 a leopard occupied 
the upper reaches of the Malchepa (Tybga mountain). 

At the end of the nineteenth and early in the twentieth century in the 


region described above leopards were subjected to intense destruction by 
every possible means, including poisoning with strychnine. The destruction 
of ungulates which serve as food for leopards also played a significant role 
182 in the population reduction of this predator. By the beginning of World 
War I the leopard population in this region had been greatly reduced and 
in other regions totally eliminated. For example, it had disappeared on the 
lower reaches of the Kishi and in most sections along the Malyi and Bol'shoi 
Laba (Nasimovich, 1941). 

In the years of the Civil War and afterward leopards were sighted once 
again in some regions where they had previously disappeared. Thus, in 1920 
they were seen on the Alousa and near Urushten camp in September, 1932 
(Olenich-Gnenenko, 1947; Burchak-Abramovich and Dzhafarov, 1949). 
Their presence was detected in the Kishi and Beloe basins from 1923 to 
1926 in the Chugush, Alamazha, and Dzhemaruk mountains; in the winter 
of 1933 to 1934 in the Malyi Laba basin; and in 1929 in the Bzyk range. 
From 1934 to 1936 this predator was sighted several times on the Chugush 
and Assari; tracks were discovered twice in Febraary, 1934, along the middle 
Berezova in Chugush region, and in February, 1936, along the Kishi River 
at Senii camp (1,400 m above sea level) (Nasimovich, 1941). 

Leopards reportedly lived in the environs of the Black Sea in the past. 
We can only add here that at the end of the nineteenth century they were 
not rare in the neighborhood of Veselii village (near Adler); they were also 
met with in the Bzyk valley and in the Pontic mountains (Dinnik, 1914). 
During the last 40 years on the Black Sea side of the Great Caucasus range, 
leopards have been noted in 1926 between the Ююsta and Agura Rivers, 
4 km from the Black Sea coast; in 1928 at Azhek village (Sochi River); 
in 1929 two animals caught around Orekhova Poly ana (Sochi River). 
Leopards were seen often in the upper reaches of the Khosta from 1927 
to 1929 and also in the upper reaches of the Sochi in legosh and Ashako 
ranges; in the summer of 1934 they were sighted in the Khostinsk branch 
of the Caucasus preserve. Finally, they occurred in the Tuapse region 
(Il'minovka village and Yakornaya Shchel' River). In 1937 leopards were 
noticed in the territory of the western section of the Caucasus preserve. 
In 1938 they were encountered a few times in the high mountains on the 
southern slope of the Great Caucasus range. In 1939 a few appeared within 
the precincts of Caucasus preserve (Ryabov, 1959). In that same year turs 
which had fallen prey to leopards were discovered in the upper reaches 
of the Malyi Laba; the predators still persisted in the eariy 1940's to the 
Dzhumunik and Dzhugush mountains (Burchak-Abramovich and Dzhafarov, 
1949; Ryabov, 1950). From the end of the 1940's and through 1957 inclusive, 
leopards were not met with in the Caucasus preserve and apparently had 
disappeared there (Ryabov, 1959). 


Leopards were observed on the upper Beloe in 1944, on Chugush 
mountain in August, 1946, and one persisted for quite some time in 1950 
around Babuk-aul village in the Lazarev region. In the autumn of 1952 
an animal was noted along the Biryuchka River in the Adlerov region. 
Instances of leopard intrusion into Lazarevskii village near Sochi (Lovetskii, 
1956), and in the autumn, 1956, on the upper Bzyk (Ryabov, 1959) have 
been described. 

Therefore, in the western Caucasus in the 1930's leopards were still 
seen occasionally primarily in two regions — the upper Kishi and Beloe — 
in the Atamazhi, Tybchi, Dzhamurk, Chugush, and Assara ranges, and the 
upper Golovinki, Bzyk, Sochi, and Khosty. A few animals lived, apparently, 
in each region. Later, they became extremely rare here.^^'^ 

In the last century in the upper part of the Kuban, leopards were 
frequently killed in the upper reaches of the Urap and Teberda. They have 
not been noted in recent decades in the Teberda preserve but some were 
caught in 1958 more south-easterly, in the Kabardino Autonomous Soviet 
Socialist Republic, on the upper Chegem, a tributary of the Terek (Alekseev, 
1958). In the former Terek and Dagestan districts leopards have been reported 

184 Fig. 102. Habitat of leopards in central Kopet-Dag. Biotope of goat (Capra 

aegagrus — left slope) and mountain sheep {Ovis amnion cycloceros — more gentle 
slopes on right). May, 1963. Photograph by N.I. Ishadov. 

^^^However. it was later reported that stray encounters at Babuk-aul (forest preserve) 
have taken place (Aleksandrov, 1965). (V.H.) 


at many places but comparatively rarely, being common only in the upper 
Avai^k Koisa. One leopard was caught not far from Evdokimov fortress 
on the Argun River (Dinnik, 1914). In the spring of 1924 another was killed 
at Agach-aul near Makhachkala city (Burchak-Abramovich and Dzhafarov, 
1949). In Dagestan at the beginning of the present century the leopard was 
encountered more or less regularly only in the region of the upper Andiisk 
and Avarsk Koisa; the animal was apparently already absent in the central 
Caucasus (Heptner and Formozov, 1941). 
183 On the southern slopes of the central parts of the Great Caucasus range 

leopard was earlier generally common in the Alazan valley, and was killed 
many times around Zakatal, Lagodekh, Nukhu, and the Bumsk gorge. In 
December, 1931 one was killed in the Zakatal'sk preserve, and in November, 
1937 at the pass opposite the city of Zakatal (Markov and Mlokosevich, 
1935; Vereshchagin, 1942). In recent years none have been seen in the 
Zakatal'sk and Lagodekh preserves, but tracks were encountered in 1937 
to the east, on the upper Tala-chai. 

All the information given above suggests that leopards are now met 
with extremely rarely in the Caucasus proper. 

In the western Trans-Caucasus and also in the coastal region reports 
of leopards were very rare even in the last century. Some animals were 
noted between Kur' and Rioni. In the Borzhomi region three were bagged 
over several years. Near Tiflis [Tblisi] one was killed in the early 1850's 
and another caught in the 1950's (A.A. Nasimovich). L-eopards were also 
not rare in the Karabakh and Talysh ranges (Radde, 1899; Dinnik, 1914). 
In the Talysh, for example, G. Radde was given 12 fresh skins in 7 weeks 
in 1866. 

Over the last 40 years in the Trans-Caucasus leopards have become 
extremely rare, and occur irregularly. They were still encountered fairly 
regularly in Talysh and Zangezur ranges, although already rare. In this period 
they were noted at the following places: 1929 (or 1930) near Kharmandal 
village on the Mugana (dead female; Sokolov, 1931); 1939 between 
Shangolu and Karabakhlyar (Sarai-Bulagsk range); 1940's in Krasnovo 
Kirovobad region; 1946 in Kul'badzhar region; 1946 in the Zangezur range 
east of the city of Ordubad (two leopards killed); 1947 in Shikh-Yurda region 
on the western slope of the Zangezur range; and 1947 on Ilanludag mountain 
and sometimes Darrydag and Megrinsk mountains (Burchak-Abramovich 
and Dzhafarov, 1949). In 1930 a leopard was caught at Ogra in the Lenkoran 
region and another animal killed there on March 13, 1930 (Starchikhin, 
1930); in 1944 a leopard was caught 30 km from Lenkoran and in 1946 
a juvenile was killed on the Talysh (Alekperov, 1947). In 1952 an animal 
was caught in the hills near Araks gorge at Neram station (Sadykov, 1952). 
In 1953 a leopard was killed in the Zangezur range near Bartsruni in the 


Azizbekov region, and in the Ararat valley near Dvin village in the Artashats 
region; in 1956 one was caught roughly 25 km east of Araks in the Vedinsk 
region in Khosrov forest where it had lived for several months (Geilikman, 
1956). In recent years leopards have been encountered from time to time 
in southern Armenia in the bCafan region. In 1957 two leopards were caught 
in Armenia, and another in the winter of 1959 (Zaritap village in the Azizbekov 
region; Mnatsakanyan, 1959). In southern Armenia in the 1950's on the 
average one leopard was killed every one to two years (Gambafyan, 1957). 
In 1964 one animal was confined to the Shams had insk region (Kasumov, 
1964). In the mid-1960 's in Annenia leopards were still encountered in the 
Zangezur range but were very rare; the population there was supplemented 
by animals coming from Iran. In the rest of the Republic this species was 
exceptionally rare and reported only as an intruder (V.G. Heptner). 

In Georgia at Akhaltsikhe a hunter killed three leopards in the winter 
of 1959 to 1960 (Galandyan, 1960). The animals had evidently arrived from 

In Azerbaidzhan in 1958 a leopard was caught in the Gaidag gorge 
(Khromov, 1958) and a second killed near Lenkoran in the winter of 1958/59. 

185 Fig. 103. Steppe -covered plateau in the higii mountains of the central Kopet-Dag — 

pasture of mountain sheep {Ovis amnion cycloceros) and site of residence and 
hunting of leopards. In the center is an old juniper tree (arborescent juniper shnib), 
a shady refuge for sheep and leopard. April, 1963. Photograph by N.I. Ishadov. 


In December, 1959 one was killed near Atrek village, Mardakerts region 
of Nagorno-Karabakh Autonomous District. In this republic one of the last 
leopards was killed in January, 1967 in the Gadruts region near Tug village 

To what extent leopards are rare in the Caucasus and in Trans-Caucasus 
can be judged from the fact that skins come singly for tanning. For example, 
only one skin was prepared in Azerbaidzhan in 1937 and 1938. In the 
preceding decade for the whole of the Caucasus, only 10 leopards were 
caught, most of which were killed in the Zangezur range and on the Talysh 
(Vereshchagin, 1947). In 1960 on the Talysh four hunters bagged only five 
184 leopards and a minimum of nine were killed throughout Azerbaidzhan from 
1960 through 1964 (A. A. Sludskii). Only two to four animals have been 
killed per annum in this republic since then (Aliev and Nasibov, 1966). 

In the Trans-Caucasus leopards were encountered from time to time 
even on the plains, especially on the Kurinsk lowlands; one was killed on 
the steppe close to Salyany (Radde, 1899). On February 20, 1946, on the 
northern Apsheron Peninsula at Bil'gya village, 25 km north of Baku, an 
animal arriving at the time of a snowstorm fell into a well (Dzhafarov, 1946; 
Alekperov, 1947). 

In the Caucasus and Trans-Caucasus leopards disappeared rapidly as 
a result of ruthless killing, cultivation, and a reduction in population of 
ungulates. In Armenia mountain goats and sheep, which live in association 
with leopards, were almost extinct even in the 1960's (V.G. Heptner). 
Leopards will evidently disappear in the Caucasus in the near future. 

Early in the present century leopards were "quite common" at Kopet- 
Dag in Turkmenia. "In 1911 1 heard about panthers everywhere and almost 
daily saw their tracks and lairs in the sands of high montane streams and 
also spoor" (Dinnik, 1914). In the first two decades of the twentieth century 
in Ashkhabad there were hunters who had each killed several leopards (V.G. 
Heptner). In the 1920's it was thought that the average density was 0.25 
animals per 100 km^ throughout the whole of the range in this republic. 
Even in the 1940's and 1950's, when mountain sheep were abundant 
everywhere in the Kopet-Dag, especially in the higher regions, leopards 
were common throughout the range in spite of a lower population than in 
the 1920's. In the Kopet-Dag some 15 leopards were killed annually in 
1940 and 1941 (V.G. Heptner). Leopards were very common in the 
mountains between Kushka and Murgab (Chengurek mountains). They were 
particularly numerous west of Badkhyz in the Gyaz'-Gyadyk mountains, 
an area extremely rich in argali sheep in those years. Leopards were regularly 
found and killed there, at the rate of 14 animals per year in 1947 and 1948, 
in a very small area (about 500 km^). The leopard population in the Gyaz'- 
Gyadyk was larger than that found anywhere else in the Soviet Union. Apart 


186 Fig. 104. Broad valley with a spring, middle belt of the central Kopet-Dag, with 

juniper, mountain maple, honeysuckle, small fruit cherry, and other vegetation, 
used as a water hole by mountain sheep (Ovis amnion cycloceros) and goats (Capra 
aegagrus), and a source of water and prey for leopard. August, 1963. Photograph 

by N.I. Ishadov. 

from the abundance of argali sheep, this was due to the arrival of migrant 
animals from Iran. In the 1960's the population shrank drastically, in part 
due to a reduction in number of mountain sheep (V.G. Heptner). 
185 Leopards were common in the Great Balkhan but due to ruthless killing 

and a sharp reduction in sheep they disappeared or became very rare in 
the 1950's. They were always rare in the Little Balkhan and long absent 
in the Kyuren'-Dag, where they probably had disappeared even in the last 
century (V.G. Heptner). In Turkmenia 19 (1959*) to 41 (1956) skins were 
tanned annually in the 1950's.^°^ 

Leopards were very rare in southwestern Tadzhikistan and southern 
Uzbekistan (Babatag, Aktau, Karatau ranges, and others). 

In southern and southeastern Trans-Baikal in the last as well as the 
present century, leopards were sighted rarely and very rarely intruded into 
the central Amur region. 

*Listed in ascending order of skin tanned — Suppl. Scient. Edit, 
^'^'^hese skins might also include those of cheetah. Some leopard skins were no doubt 
retained by hunters. 


In the last century in the Far East, in the Ussuri territory leopards were 
rare. They were found often only in regions adjoining China. In 1867 during 
a six-month journey throughout the Ussuri territory, covering a total of 2,250 
km (partly by boat), N.M. Przheval'skii (1870) discovered only two sets 
of leopard tracks and obtained a single skin with much difficulty. Much 
of his route lay along the coast of the Sea of Japan, covering places where 
leopards were reportedly often encountered. 

Leopards were rare in the Far East even in the early years of the present 
century. In 1912 two animals were killed in the Amur region, and eleven in 
the Primorsk district (Baikov, 1927a). According to other data, however, only 
one or two were caught annually in these regions between 1911 and 1914 
(Tselishchev, 1924). In the 1920's two or three animals were caught per 
year in the Ussuri territory (Solov'ev, 1925). Leopards were even rarer in 
the 1950's and 1960's. However, encounters were more frequent due to 
inteasive cultivation of formeriy virgin areas and human population growth. 
As in the last century, this predator is now "not a rarity" along the western 
coast of the Amur Gulf. Some 15 to 20 leopards presently live in the montane 
forest region lying to the north of Suifun River and thence southwest to the 
186 state frontiers with the Korean People's Democratic Republic and the 
Chinese People's Republic (Vasil'ev et al., 1965). According to other data 
this number represents the leopard population inhabiting the whole of the 
Primor'e (G.F. Bromlei).^^^^ 

In "Kedrovaya Pad' " preserve (southwest of Vladivostok) four to six 
leopards were regularly seen in an area of 9,500 hectares in 1940 and 1941 
(Mirolyubov, 1941); a family of two to four animals lived there in a section 
40 km^ in the mid-1960's (Vasil'ev et al., 1965). The high population 
density of leopards in this preserve is explained by the abundance of 
ungulates there. 

Habitat. In the Cauciisus leopards inhabit subalpine steppe-covered 
meadows, deciduous forests, and dense shrubs; they are generally confined 
to rock outcrops and talus slopes. The main conditions determining their 
habitat are a high population of bezoar goats, turs, roe deer, chamois, deer, 
and wild pigs, and the availability of sections with shallow snow in winter. 
Thus, in the western Caucasus leopards live high in the hills and are seen 
quite often in summer in the alpine zone and even at permanent snow line 
at a height of 2,600 to 2,700 m above sea level, and sometimes even higher; 
in the central part of the range they ascend to 3,000 to 3,500 m above sea 
level (Dinnik, 1914). They are confined in the latter area to a fairly high 
level even in winter and have been sighted at 1,500 m (February, 1934; 
Berezova River) and 1,400 m above sea level (Febmary, 1936; Kishi River). 

^^In 1970, 32 to 35 leopards were counted in Ussuri territory (S.P. Kucherenko). (V.H.) 


187 Fig. 105. Steppe-covered plateau with juniper trees (arbore.scent juniper siiruLxs) 

and limestone cliffs at the top of the Great Balkhan mountains, a place formerly 

very rich in mountain sheep (Ovis amnion cycloceros — plateau) and goats {Capra 

aegagrus — cliffs). Leopards were also common there. About 2,000 m above sea 

level. February, 1963. Photograph by E.N. Matyushkin. 

On both occasions they were close to the wintering sites of chamois. The 
depth of the snow cover ranged from 70 to 100 cm (A. A. Nasimovich). These 
mountains are rich in turs, chamois, and at times deer. On the Black Sea 
side of the western Caucasus leopards often do not reach the alpine zone, 
characterized by extreme rockiness, slopes overgrown with dense cherry 
laurel, azalea, Pontic rhododendron, and Caucasian bilberry. Many areas 
of these mountains are almost impenetrable to human. Chamois are quite 
common in the higher levels and roe deer and wild pigs also occur at sui- 
188 table places. In these mountains leopards range from the sea coast up to 
the mountain tops. 

In the Trans-Caucasus this predator lives (lived) in the treeless or nearly 
treeless rocky mountains and gorges where bezoar goats and Traas-Caucasian 
mountain sheep are abundant. In southern Armenia it is confined to 550 
to 950 m above sea level on slopes with steppe vegetation and rarely ventures 
into forest regions (Dal', 1954). Leopards sometimes live even in the plains, 
for example in the Lenkoran lowlands, if they are covered with dense forests 
or reeds, prickly shrubs, and various kinds of lianas, and inhabited by wild 


187 Fig. 106. Gorge dissecting a plateau summit. Left, an old juniper tree (arborescent 

juniper shrub). Typical leopard habitat in the Great Balkhan mountains. February, 
1963. Photograph by E.N. Matyushkin. 

pigs. On the level steppes leopards occur only as intruders. In the Kurinsk 
lowlands they are encountered from time to time in reed-cattail thickets 
around water reservoirs, and also in tugais along the Kur' and Araks (N.K. 

In Turkmenia leopards mainly inhabit mountains featuring rocks and 
gorges, and almost devoid of trees, or sparsely covered with juniper and 
pistachio. Along the cliff slopes of the gorges of these mountains and uplands, 
tangles of dog rose, blackberry, barberry, and in places native pistachio, 
are encountered, which grow on the terraces, "shelves", and in their bottoms. 
Leopards are confined here to sites rich in bezoar goats and Turkmenian 
mountain sheep {Ovis amnion cycloceros) or only the latter; the predator 
ascends in summer to the uppermost levels and descends in winter to lower 
levels following goats and sheep. 

The Gyaz'-Gyadyk mountains in southem Turkmenia (Badkhyz), where 
leopards are still relatively common, are not high, but are interrupted 
by deep gorges with steep slopes, and limestone and sandstone outcrops. 


188 Fig. 107. Slopes and small valley with pistachio thickets and rich herbaceous cover 

of sagebioish, meadow grass, and giant umbellifers in Gyaz'-Gyadyk. Characteristic 

biotope of argali sheep (Ovis amnion cycloceros), and leopards found in association 

with them. Pistachio represents a favorite shady refuge of both species. Badkhyz 

preseive. October, 1959. Photograph by A.A. Sludskii. 

Mountain slopes are covered with sedges, meadow grass, and forbs while 
along the range, its slopes to the very bottom are covered with perennial 
trees of the true pistachio, the diameter of which at ground level may reach 
1.0 m. Sometimes these trees form characteristic bushes. They are well 
separated and do not form close clusters ("Pistachio savanna"). Even old 
trees are not tall but very dense; their crowns form a semicircular dark green 
marquee since their low branches almost reach the ground. 
189 At the end of summer, when pistachio fruits ripen, ungulates congregate 

there; wild pigs, Turkmenian sheep, goitered gazelles, and porcupines. 
Leopards are attracted by the abundance of prey. Goitered gazelles are 
particularly numerous around pistachio trees in arid years. Along the bottom 
of gorges, usually overgrown with wild figs, small saline springs occur, 
for example at Kerlek, containing 8.0 to 20.0 g salts per liter. Reeds grow in 
a narrow belt along some of these springs and wild pigs inhabit them. Kulan, 
wild sheep, sometimes goitered gazelle, and various carnivores — wolf, 
hyaena, and cats — gather at the more accessible springs in summer. Leopards 
also drink from such springs. Predators lie in wait for ungulates at these 
watering places. In the Badkhyz leopards have been found in the deep Er- 



189 Fig. 108. Spring in one of the gorges of tiie Gyaz'-Gyadyk mountains serving 

as a watering site and hunting ground for leopards. Slope streaked with trails of 

ungulates coming to water, mainly mountain sheep, also wild pigs, .sometimes 

goitered gazelles and kulans. Badkhyz preserve, southern Turkmenia. February, 

1965. Photograph by V.G. Heptner. 

oilan-duz depression, hunting for wild sheep among chinks [arroyos] over- 
grown with herbs and shrubs such as sahwort, locoweed, and others. Along 
these cliffs sheep from Gyaz'-Gyadyk, trailed by leopards, enter the very 
heart of the desert at Dyzyl-Dzhar where, in addition to sheep, goitered 
gazelle, some wild pigs, and kulan are common visitors (Heptner, 1956; A. A. 
Sludskii). In the Atrek basin in Kopet-Dag (Sumbar and Chandyr Rivers), 


190 Fig. 109. Spring at site of Kerlek, watering place of leopards in Gyaz'-Gyadyk 

mountains. Badkhyz preserve, southern Turkmenia. May, 1962. Photograph by 

A. A. Sludskii. 

192 leopards also live in large gorges with giant nut-bearing trees, thickets of 
fig, and other trees entwined with vines, impenetrable tangles of blackberry, 
etc. (Ai-Dere, Pop-Dere, etc.) (V.G. Heptner). 

The leopard's range in Turkmenia almost totally coincides with that 
of Turkmenian sheep. This carnivore is quite common in regions where 
bezoar goats are absent or rarely encountered; it is also absent at places 
where goitered gazelle are numerous, but is present even in unfavorable 
deserts if they are inhabited by sheep (Badkhyz). On moving into a desert 
the carnivore confines itself to cliffe where it can find shelter (Heptner, 1956). 

In Tadzhikistan and Turkmenia leopards live in low mountains covered 
with desert vegetation and normally do not ascend higher than 1,500 m 
above sea level. In Babatag (Uzbekistan) leopards keep to deep and narrow 
gorges and precipitous rocky slopes which alternate with relatively gentle 
slopes covered with pistachio shrubs. At the bottom of these gorges many 
fresh or saline springs are present, and tamarisk, poplar, and sometimes reeds, 
ambary, and grapevines are relatively abundant. Ironwood, almond bushes, 
and Persian ironwood grow on the steep rocky slopes near such places. 
Mountain sheep and markhor, the main prey of leopards, live on the slopes 
of these gorges and in the watersheds between them. Leopards in Babatag 


190 Fig. 110. Saline laices, buttes, and chinlc wall (right — habitat of mountain sheep 

and leopards. Hyaena also live here. Er-oilan-duz depression in Badkhyz desert, 
southern Turkmenia. May, 1962. Photograph by A.A. Sludskii. 

live at heights of 600 to 900 m above sea level (Bogdanov, 1952; Ishunin, 

In the Far East, in Primor'e, leopards are confined to a narrow coastal 
strip with Manchurian type broad-leaved forests and at places with a rather 
low snow cover. They visit rocky sections along the sea coast very rarely, 
on the average two or three days in a year. For example, on Tuman mountain 
leopards were sighted only in March, 1942; December, 1948 [sic]; and 
January, 1946. Between 1944 and 1946 some stray tracks of leopards were 
detected on two occasions in autumn along the coast extending from 
Valentine Bay to Preobrazheniya Bay (Bromlei, 1963). The animal has been 
sighted on occasion, mainly in winter, deep inside the mountains, on their 
western slopes. Evidently the high snow cover of these areas poses a serious 
obstacle to movement. On Cape Gamov, when the snow cover is high, 
leopards find movement very difficult and prefer confinement to one place 
193 on talus slopes (V.V., 1927). In winter, during wanderings, they keep to 
snowfree rocky slopes facing south. Rocky sections are favored over adjacent 
sections devoid of rocks. This carnivore generally avoids roads, human and 
animal trails, frozen ice crusts, and rivers; however, it takes advantage of 
deer trails for hunting in certain areas. 


191 Fig. 111. Buttes of the Ei"-oilan4liiz depression; nortliem chink visible in background 

and bottom covered with solonetzes and saxaul shrubs. Habitat of argali sheep, 
goitered gazelle, kulan, leopard, and hyaena. June, 1948. Photograph by V.G. Heptner. 

191 Fig. 112. Gigantic argillaceous cliffs ("sheep cliffs") in Kyzyl-Dzhar gorge 

dissecting Badkhyz desert plateau. Habitat of arkhar sheep, visited by leopards. 
Southern Turkmenia. February, 1961. Photograph by M.V. Heptner. 


192 Fig. 113. Forest of "Kedrovaya Pad' " preserve. Site of permanent residence of 

leopards. Tiger, leopard cat, sika deer, etc. also live here. June, 1958. Photograph 
by A.G. Pankrat'ev. 

In the Far East leopards live not only in montane forest regions but 
also in valleys and plains. For example, at the end of the nineteenth century, 
these animals were met with in meadowlands in the Pre-Khanka lowland. 
Nowadays none are found in the lowland meadow-forest sections of the 
Mo and Lefu interfluve ([Lake] Khanka tributaries) of the Pre-Khanka 
lowland. South of Lake Khanka in 1956 to 1961 these predators lived in 
rocky creeks and among reeds in the floodplains of the Chapigos valley. 
In the 1950's they were seen from time to time in meadow-forest sections 
in the floodplains of the Amur and Bir (Rakov, 1965). Their distribution 
in the Far East is usually associated with the presence of sika and roe deer. 
It is quite possible that in Primor'e this species is only an intruder and hence 
breeding there would be merely incidental (G.F. Bromlei). 

Food. Wild ungulates — bezoar goats, turs, mountain sheep, chamois, 
roe deer, deer, and wild pigs — constitute the main prey of leopards in the 
Caucasus and Trans-Caucasus. Sometimes they also catch European hare, 
pheasant, rock partridge, Caucasian black grouse, snowcock, and porcupines 
(in Talysh) (Dinnik, 1914; Vereshchagin, 1942). At places where ungulates 
are abundant, for example in the former Kuban district, leopards almost do 


193 Fig. 114. Deer Cliff in "Kedrovaya Pad' '' preserve near Vladivostok. Regular 

favorite habitat of leopards. June, 1964. Photograph by A.G. Pankrat'ev. 

not attack domestic animals, but in the Trans-Caucasus attacks on various 
domestic animals, including dogs and poultry, have been reported frequently. 
Bones of horses, asses, cattle, wild goats, and birds have been found around 
a leopard's lair in the Zangezur range near Kalita village (Burchak- 
Abramovich and Dzhafarov, 1949). 

In Turkmenia, in the Kopet-Dag, leopards hunt mainly bezoar goats 
and Turkmenian mountain sheep, and more rarely wild pigs and domestic 
animals such as cattle, horses, asses, sheep, goats, and dogs. Not only the 
young but even the aduhs of these animals are attacked (Dinnik, 1914; 
194 Bil'kevich, 1924; Morits, 1935). Around the lair of a leopard in the Gyaz'- 
Gyadyk (Akar Cheshma) there were many bones of goitered gazelle 
(Filippov, 1956). The main prey of leopards are Kopet-Dag sheep in most 
regions of Turkmenia, especially in Badkhyz (Heptner, 1956). Sometimes 
leopards attack porcupines but pay the price. In the soles of one leopard's 
paw were found nine pieces of porcupine quills measuring up to 1.0 cm. 
Necrosis had set in around the pierced areas (S.I. Bil'kevich). In February, 
1951, in Badkhyz (Gyaz'-Gyadyk, Kepel village) a leopard was killed which 
had many pieces of porcupine quills in its paws (Rustamov and Shcherbina, 


1957). Another one caught on August 4, 1960, in Badkhyz had quills up 
to 4.0 cm in its limbs and other body parts (Gorelov, 1963). In March, 1955, 
a leopard was located in the burrow of a porcupine which it had killed 
(Dement 'ev and Rustamov, 1956). 

Leopards take to attacking domestic cattle mainly in snowy winters 
or when the population of wild ungulates has shrunk. For example, in the 
extremely snowy winter of 1921 an animal lived quite close to a village 
and daily stole a sheep (Bil'kevich, 1924). The reduction in number of bezoar 
goats and Turkmenian sheep observed in recent decades in Kopet-Dag has 

195 Fig. 115. Leopard habitat in "Kedrovaya Pad' " preserve, near Vladivostok. 

August, 1969. Photograph by G.N. Simkin. 


led to an increase in attacks by leopards on domestic animals in that region. 

195 Some feed on cattle fairly regularly, and are termed "village panthers" in 
India. As late as the spring of 1951 in the Kyzyl-Arvats region three leopards 
were killed when they attacked cattle. One animal regularly attacked horses 
and killed 9 in 1949, 17 in 1950, and 3 in the first three months of 1951. To 
safeguard them against leopard attacks horses in a collective farm had to be 
driven to another region. A leopard killed a hoise even in the Badkhyz preserve 
(Dement 'ev and Rustamov, 1956). In Dmitrievka village, 50 km from Goek- 
Tepe, the predator caught nearly every dog available (Dinnik, 1914). 

In the Ussuri territory the main prey of leopards are roe deer, sika deer, 
Manchurian wapiti, musk deer, moose, and wild pigs; more rarely the 
predator catches hare, badger, fowl, and even "mice" (Baikov, 1915 and 
1927; Yankovskii, 1882). In "Kedrovaya Pad' " preserve (southwest of 
Vladivostok) roe deer is the main prey of leopards year-round (Vasil'ev 
et al., 1965). In the southern Ussuri territory (Tumannaya knoll) it catches 
hares and from time to time gorals (G.F. Bromlei). In "Kedrovaya Pad' " 
preserve leopards attack raccoon dogs, often abandoning them; they prey on 
young, Eurasian black bears (Ursiis tibetanus) less than two years old 
(Vasil'ev et al., 1965). These bears are preyed upon even in the north, in the 
Pri-Amur (Sysoev, 1966). In Primor'e, leopards attack sika deer held in parks 
(Menard, 1930; G.F. Bromlei). In the forest nursery at Pushkino a leopard 

196 killed in one day three deer, two the next day, and so on. Because of the 
extreme rarity of leopards in Sudzukhin preserve, between 1936 and 1948 
not a single mortality of sika deer was attributable to this predator (Bromlei, 

The stomach of a large male caught on August 4, 1960 at Badkhyz 
contained 2.0 kg of meat from a mountain sheep (Gorelov, 1963). In 
zoological gardens leopards are fed daily about 3.0 kg of meat (Obudhova 
and Shakhnazarov, 1949). The report that a hungry predator can consume 
almost an entire roe deer in a single day (Baikov, 1927) is clearly an 

In China leopards catch deer, wild pigs, gorals, monkeys, hares, large 
birds, and domestic animals, particularly dogs and pigs (Shou, 1958). In 
India this predator hunts for sambar deer, wild pigs, jackals, monkeys, hares, 
and porcupines, and also eats lizards, snakes, and crabs. In Kashmir it attacks 
hangul deer when their movement is impeded by deep snow. Leopards often 
prowl near villages and sometimes become a pest of domestic cattle, 
destroying calves, asses, horses, goats, sheep, and other animals, but most 
often dogs, for which they will even break into a house. Jackals and dogs 
represent their favorite prey. They also eat carrion irrespective of the degree 
of decomposition. In Africa the biggest animals attacked by leopards are 
camels, cows, kudu antelope, and asses. The predator usually feeds on 


f * - fc .. 

195 Fig. 116. Site of transient and resident leopards in Sudzukliin preserve on the eastern 

slope of Ta-Chindzhan range. Southern Ussuri territory. August, 1958. Photograph 
by V.E. Prisyazhnyuk. 

moderate-sized and small animals such as small antelopes, wart hogs, sheep, 
goats, pigs, aardvarks, monkeys (baboons), porcupines, domestic fowl, and 
small rodents; it also attacks jackals, foxes, genets, and domestic dogs. 
Leopards sometimes attack man and some become man-eaters, i.e., man 
may serve as their main food item (see "Practical Significance"). 

Home range. Not much is known about this. Zoologists are of the opinion 
that, outside the breeding season, leopards prowl continually in search of sites 


196 Fig. 117. Montane mixed broad-leaved forest witli an admixture of fir and Korean 

nut-pines. Habitat of leopards in "Kedrovaya Pad' " preserve near Vladivostok. 
December, 1962. Photograph by A.G. Pankrat'ev. 

abundant in wild animals. This view is obviously only partly justified. At 
places where wild animals are abundant leopards live permanently or perform 
only vertical migrations, trailing herds of ungulates and avoiding snow. A 
family of two to four leopards lives regularly in "Kedrovaya Pad' " preserve 
in Primor'e. Year after year they have occupied the same section and rarely 
197 gone outside the preserve boundaries. This section covers the upper reaches 
of the Kedrovaya River where forests are dense and narrow mountain crests 
and peaks with rocks and talus slopes stand out prominently. The area of 
this section is about 40 km^ and the leopards have made permanent trails 
within its confines (Vasil'ev et al., 1965). 

Two or three leopards lived permanently in the Chapigou River valley 
of Primor'e from 1956 through 1961 and possibly constituted a single family. 
These animals were seen in an oval-shaped hunting territory, the longer side 
of which was about 40 km. Annually, after the commencement of frosts, the 
predators descended from talus slopes into valley sections of reeds housing 
abundant roe deer and raccoon dogs. In 1960 and 1961 a lone leopard on 
the southern slope of the Pidan range occupied a hunting region 30 to 


197 Fig. 118. Mixed coniferous — broad-leaved forest in "Kedrovaya Pad' "" preserve 

near Vladivostok. Habitat of leopards. Spiny actinidia vine on Korean nut-pine 
visible in foreground. October, 1965. Photograph by A.G. Pankrat'ev. 

35 km long and 10 to 15 km wide and fed mainly on roe deer (Rakov, 
1956). A leopard remained close to a deer park for several years and 
eventually had to be put down (Menard, 1930). 

In the Caucasus in recent decades the individual territories of leopards 
were very large, occupying as much as 100 km^ (Nasimovich, 1952). 

Burrows and shelters. The den in which a leopard raises her cubs is 
set up in a cave under an overhanging rock or under a wind-felled tree. 



located usually in a particularly dense, inaccessible section of a gorge. A 
lair in the Kishi valley (western Caucasus) was built on the ground in a 
wide patch of uprooted trees. The ground at the site of the den and around 
it had been churned and the grass well beaten. Feces of the female and 
cubs and bones of various ungulates were scattered around the den. A second 
den in the same patch of wind-felled trees had been set up under a rock 
(Dinnik, 1914). In Turkmenia a leopard was caught in the burrow of a 
porcupine (see above). A female with two cubs was found in the burrow 
of this rodent near Akar-Cheshma. Around the burrow were many bones 
of goitered gazelle (Filippov, 1936). In northeastern China shelters for raising 
cubs are located among dense shrubs, near talus slopes on the southern 
slopes of hills, in caves, and under overhanging rocks. The den [floor] is 
covered with dry leaves and grass prior to parturition (Baikov, 1915). If 
a den with very small cubs is threatened, the mother carries them in her 
teeth, one at a time, to a new site. 

Males and lone females also rest during the day in caves, rock fissures, 
or dense thickets. In Turkmenia caves and very deep niches in rocks usually 
serve as leopard shelters (V.G. Heptner). A leopard shelters on warm summer 
days in the shade of an overhanging rock but loves to sun itself on an exposed 
rock in cold weather. Sometimes the predator sleeps during the day in well- 


', '" i^'bt' 

Fig. 119. Site of permanent occupation of leopards in "K.edrovaya Pad' "' preserve, 
near Vladivostok. September, 1970. Photograph by V.G. Heptner. 


protected places, such as open sites on slopes, behind rocks on ledges, etc. 
(V.G. Heptner). 

In India, as well as in Turkestan, the den is sometimes made in a 
porcupine burrow (Pocock, 1939). 

Daily activity and behavior. Leopards set out to hunt usually an hour 
or two before sunset, hunting through the first half of the night, and again 
in early morning. In spite of the fact that a leopard is essentially a crepu-scular 
animal, from time to time it chases prey even during the day, especially 
on cloudy, cold days and in winter. It goes to water holes with the onset 
of twilight, but sometimes even during the day. 

The leopard is a cautious, secretive animal, but never timid. On 
encountering a man, it does not attempt to hide quickly as do many animals, 
but withdraws unhurriedly and with no exhibition of fear. Sometimes it does 
not run away even when a shot has misfired. Instances are known when 
leopards, in sight of hunters, carried off a goat wounded by them, or of 
pursuing game unmindful of the shouts of the people or their shots. Unlike 
ounce and cheetah, an injured leopard often attacks the hunter or chases 

198 Fig. 120. Broad-leaved forest with an admixture of nut pines and fir. Biotope of 

leopards. "Kedrovaya Pad' " preserve, near Vladivostok. September, 1970. 
Photograph by V.G. Heptner. 


him. Hence in most southern Asian countries leopard hunting is considered 
extremely dangerous. Elsewhere, instances have been recorded of an animal 
wounding several men who set out to catch the predator. When attacking 
a man a leopard rears up on its hind legs and strikes him with its forepaws 
or attempts to grab him by the throat. 

A leopard can conceal itself exceedingly well and often remain within 
just 10 m of men passing by. It climbs well and in rescuing itself from pursuit 
generally climbs inaccessible rocks or trees and conceals itself, making 
detection very difficult. Its every movement is graceful, supple, and agile. 
When a leopard moves, it progresses as quietly and lightly as a small cat. 

This predator usually avoids water and prefers to cross small brooks 
on fallen trees or a bridge. On occasion it goes into water and swims across 
large rivers. In the western Caucasus in February one year a leopard crossed 
a fiord through an unfrozen rivulet, the depth of which was 50 cm (A.A. 

Vision and audition are excellent in leopards but olfaction poor, due 
to which these animals do not hunt by scent. They employ various methods 
for hunting. Usually the predator will lie in wait for prey, hiding in bushes 
among rocks, on an outcrop or tree near a trail, water hole, or solonetzes 
and attack the unsuspecting victim from an ambush in a single leap of 6.0 
to 8.0 m. On spotting the prey the predator conceals itself and crawls forward 
so quietly that detection is extremely difficult. In winter, on a snowfree slope 
with a background of dried leaves or grass, it is almost impossible to locate 
a concealed leopard even at a distance of 50 m. In grass 50 cm high, when 
the predator starts to creep, it completely disappears from view. When the 
quarry is close by, the leopard chases it in long bounds. Sometimes it sprints 
after the quarry, executing bounds of 7.0 to 8.0 m length so rapidly that 
it can quickly overtake a roe deer, goral, musk deer, bezoar goat, or dog. 
Such chases of deer, roe deer, and chamois have been observed in the 
Caucasus and chases of bezoar goats seen in Kopet-Dag. Leopards are usually 
capable of overtaking quan-y over a distance of not more than 40 to 50 m. 

In "Kedrovaya Pad' " preserve in Primor'e, in winter the leopard 
conceals itself to catch roe deer and creeps forward from the leeward side 
of the prey in the late morning hours when, after grazing, deer climb the 
slopes toward their beds. Sometimes a leopard catches one of them by hiding 
under coastal cliffs (Vasil'ev et al., 1965). 

In attacking large animals a leopard generally rears up on its hind legs 
and inflicts severe blows with its forepaws. In the case of small animals 
it twists the neck in one stroke and then bites the skull. Larger animals 
are first knocked down, after which the predator clamps its jaws on the neck 
at the occiput or rips open the throat. A female sheep in Kopet-Dag killed 
200 by a leopard had her throat slit, withers broken, and wide gaping wounds 


on the muzzle inflicted by the claws; the bone under the eyes was evidently 
pierced by the large claws of the forepaws (Morits, 1935). In killing a 
porcupine the predator grabs it from the front just behind the head, thus 
protecting itself from the quills (Corbett, 1957). 

On entering an area rich in game, a leopard usually kills a single animal 
and does not frighten the rest. In this respect it is similar to tiger but differs 
from wolf, which usually attempts to wound several animals and thus drives 
them away. Even within a deer park a leopard usually kills only one deer 
at a time. 

In the Ussuri territory the leopard hides the carcass of a roe deer among 
the roots of upturned trees or under wind-felled trees, feeds on it for two to 
four days, and hides nearby (Vasil'ev et al., 1965). The predator sometimes 
conceals the remains of its prey in hollows and rocks, or in the tropics, in trees. 

In Kopet-Dag the leopard usually returns to any unfinished prey. Quite 
often it rests during the day near the carcass of a killed animal and drives 
off vultures. These birds, numerous in this region, can pick clean the carcass 
of an argali sheep within an hour. Evidently to safeguard its prey, it will 
jump to a height of several meters, springing into a juniper tree with the 
quarry in its mouth. In spite of its comparatively small size, the leopard 
possesses tremendous strength. Once this predator was seen carrying a young 
horse it had killed over rocks for a distance of 50 m. On another occasion 
a leopard jumped onto a rock 3.0 m high with a mountain sheep clamped 
between its teeth. Ascending upward along a steep slope, this cat can carry 
a female bezoar goat rather effortlessly (V.G. Heptner). 

Seasonal migrations and transgressions. As noted above, outside the 
breeding season when food is inadequate or access to it difficult (deep snow 
cover, thin snow cnist), leopards migrate from one area to another. Fairly 
regular vertical migrations of this animal have been observed in the Caucasus 
where, depending on snow thaw in spring, the predator ascends to the alpine 
zone, trailing ungulates, and descends with them in autumn. Normally an 
inhabitant of the mountains, it sometimes migrates far into the plains (100 
km or more). 

In the Caucasus, in Dagestan, in the early part of the present century 
leopards, though very rare, were nonetheless sighted fairiy regulariy in the 
upper reaches of the Andiisk and Avarsk Koksa, and in the spring of 1924 
caught at Agach-aul near Makhachkala (Burchak-Abramovich and 
Dzhafarov, 1949). On the Black Sea coast leopards were sighted often at 
the end of the last century, and in the 1920's to 1930's, at Adler, Khosta, 
and Sochi. iThey were found there even later. In 1950, for example, a leopard 
resided for quite some time near Babuk-aul village in the Lazarev region 
and in the autumn of 1952 one was noted on the Biryuchka River at Lazarev 
village near Sochi (Lovetskii, 1956). 


In the Trans-Caucasus leopards were sighted fairly regularly in the 
present century in the Talysh and Zangezur ranges from where they spread 
out in different directions. Intrusions into the plains of this country possibly 
occurred from other places, for example the Great Caucasus range. On 
December 6, 1930, a dead female was found near Kharmandal on the 
Mugana (Sokolov, 1931). Even earlier, at the end of the last century, leopards 
were caught in the steppes near the city of Salyanyi (Radde, 1899). On 
February 20, 1946, a male was killed on the northern Apsheron Peninsula 
near Bil'gya, 25 km north of the city of Baku. During a severe snowstorm 
the animal entered a densely populated region and fell into a dry well 
(Dzhafarov, 1946; Alekperov, 1947). During the last century and in the 
1950 's leopards have been caught near Tbilisi (A. A. Sludskii). 

Leopards used to cross into Turkmenia from Iran and Afghanistan but 
not regularly and not at a given time of year. Information about regular 

203 Fig. 121. Leopard track in mud. Cartiidge placed for scale. Badkliyz preserve, 

southern Turkmenia. June, 1961. Photograph by Yu.K. Gorelov. 


migrations into Afghanistan (Herov, 1932) has not been substantiated. In 
this area animals from the Kopet-Dag foothills sometimes entered the plains 
but usually traveled only a few kilometers (V.G. Keptner). 
201 In the last century leopards were seen occasionally in southern and 

southwestem Trans-Baikal (Cherkasov, 1884; Baikov, 1927). One was killed 
within Trans-Baikal in 1912 (Baikov, 1927). By the 1950's their population 
has evidently increased somewhat. At the end of March, 1952, at Nizhnyaya 
Bareya of the Nerchinsk-Zavod region of Chita district, two attacked a flock 
of sheep and one was killed. Leopards intrude into Trans-Baikal from the 
Great IGiingan range of northeastern China. 

Leopards were occasional visitors on the midreaches of the Amur 
(Przheval'skii, 1870), on the Kumara River (Khumaerkhe), on the Amur- 
Zeya plateau at 51°45' N. lat. (Baikov, 1927), along the Tyrma River (left 
tributary of Bureya (Middendorff, 1867), and in the Bureinskii range (Radde, 
1862). Very exceptionally one was sighted in November, 1950 on the middle 
course and at the mouth of the Bira (left tributary of the middle course 
of the Amur in Birobidzhan). Leopards are not known to people living in 
the Soviet Little Khingan range, which would indicate that this animal rarely 
intrudes into this region (Rakov, 1965). 

203 Fig. 122. Leopard track near spring in Nardevanly settlement. Size of track 12 

(width) cm x 11 cm. Badkhyz preserve. Turkmenia. April, 1961. Photograph by 

Yu.K. Gorelov. 


In the basin of the lower course of the Ussuri leopards formerly 
penetrated to Khor (around 48° N. lat.) (Baikov, 1927). In recent decades 
none has been seen that far north and reports of more southern visits in 
this region are rare. In November, 1954, one animal was sighted in the upper 
reaches of the Pervoe Sed'moe (right tributary of lower Ussuri) about 30 
km south of Vyazem. In October, 1939, a lone leopard was killed in the 
Levyi Podkhorenok basin and in October, 1954, another sighted in the middle 
reaches of the Kanikheza (tributary of the middle Bikin) (Rakov, 1965). In 
the basin of Lake Khanka, at Beichikhe, leopard intrusions were recorded 
in January, 1957, and December, 1960. These predators usually penetrate 
there from China on the trail of migrating roe deer (Rakov, 1965). 

It has been suggested that most of the leopards sighted in the Primor'e 
and other regions of the Soviet Far East were intruders from the Korean 
Peninsula and China and did not breed within the confines of the Soviet 
Union. To date no litters of these cats have been reportexi in the USSR 
(G. F. Bromlei). In Suputin preserve they are sometimes seen in summer, 
arriving from the south, but remain for a ver)' short period before returning 
to their usual habitat (Bromlei and Gutnikova, 1955). On the other hand, 
leopards not only live regularly in "Kedrovaya Pad' " preserve (south- 
western Vladivostok) but also breed there (A.G. Pankrat'ev).- 

Reproduction. Very little is known about this process among leopards 
under natural conditions, but it has been studied well in captive animals. 

In the Caucasus these animals usually become sexually aroused in 
January (Vereshchagin, 1942), in Turkmenia, in the Kopet-Dag, in 
December-January and again at the end of November (V.G. Heptner), and 
in Primor'e ("Kedrovaya Pad' " preserve) (Vasil'ev et al., 1965) and 
northeastern China (Manchuria) in January (Baikov, 1914). Judging from 
the dates of sightings of small cubs, mating in the USSR may also occur 
in other seasons of the year. A few males may follow a single female in 
estrus and serious combats, accompanied by high-pitched cries, ensue among 
them. In Kopet-Dag groups of up to five animals have been seen (V.G. 
Heptner). A female in heat urinates frequently, rolls on her back, arches 
her back in a characteristic manner whereby the rump is raised, makes typical 
rumbling and purring sounds, and stops feeding temporarily or eats very little. 

Among animals held in zoological gardens estrus has been detected 
at various times of the year, and at intervals of every two to three months 
if the female happened to remain barren (sometimes it occurs each months 
or only twice a year). Estrus lasts 12 to 18 days and in exceptional cases 
up to 25 days. When the female is stimulated the animals mate many times 
a day and the intervals between consecutive copulations may be three to 
four minutes to a few hours (Shereshevskii, 1940). Gestation requires 90 
to 105 days, but usually 92 to 95 (Shortridge, 1934; Shereshevskii, 1940). 


204 Fig. 123. Trail of a leopard (female) which had been frightened away from her 

prey. "Kedrovaya Pad' " preserve, near Vladivostok. April, 1965. Photograph 
by A.G. Pankrat'ev. 


Parturition occurs mostly at night and the process normally requires six 
to ten hours. 

Everywhere within the USSR and also in northeastern China cubs are 
seen most often at the end of March to April and in April and May (Primor'e, 
"Kedrovaya Pad' " preserve; Vasil'ev et al., 1965) but in Kopet-Dag usually 
in May (V.G. Heptner). Sometimes cubs are found in other months however. 
Thus, on December 11, 1953, in Badkhyz preserve, in the Gyaz'-Gyadyk 
mountains (southern Turkmenia) in Nerdevanly gorge, two cubs with their 
eyes still closed were found, one of which opened its eyes only on December 
15 (Rustamov and Shcherbina, 1957; Yu.K. Gorelov). Evidently in the 


204 Fig. 124. Individual track of a leopard (female) on fine, wet snow. Match box 

for scale. "Kedrovaya Pad' " preserve. April, 1965. Photograph by A.G. Pankrat'ev. 

southern part of the leopard range, no definite season for parturition exists. 
In northern China cubs are usually seen in spring while south of Hwang 
Ho littering may occur at any time of the year. In Yunnan province cubs 
are more often sighted in spring but young have been reported in other 
seasons also. In India cubs are generally born in February or March 
(Blenford, 1888-1891*). In zoological gardens cubs appear at any time of 
year. In London Zoological Garden they appear more often at the end of 
February to early March, followed by end of April to May, June and July, 
end of September to October to the first half of November, and from the 
second half of December (Tsukerman, 1953**). 

The number of cubs in a litter in the southern part of the range and 
in zoological gardens varies from one to four, and in exceptional cases 
reaches six. In China a litter consists of one to five cubs, in the northern 
provinces more often three or four, and in the southern (Yunnan) usually 
two. In the wilds a lone cub is encountered very rarely. In eastern Asia a 
litter usually consists of two cubs (Tate, 1947). The number of young in 
a litter within the Soviet Union has not been established for want of adequate 
data, but most often consists of two or three. In the western Caucasus a 

^[5/c]; Blanford. Not in Literature Cited — Sci. Ed. 
"*[5/c]; Zuckerman. Not in Literature Cited — Sci. Ed. 


female with two cubs was caught (Dinnik, 1914). In the Geok-Tepi region 
of Turkmenia a female with two young was sighted. In Badkhyz two litters 
with two cubs each were found (Filippov, 1936). In Februar}', 1959, near 
Ashkhabad a female with a single cub was caught (Polozov, 1959) and in 
the same region in December another with two cubs trailing behind her. 
Families consisting of three members have been sighted time and again in 
Turkmenia (Dement 'ev and Rustamov, 1956). The number of cubs there 
is usually two but may reach four (V.G. Heptner). In Primor'e, in 
"Kedrovaya Pad' ", litters comprise one to three cubs (Vasil'ev et al, 1965). 
Growth, development, and molt. Cubs are bom blind and helpless but 
covered with dense and fairly long hair. The main color tone of the coat 
in the upper part of the body in newborns from the nose to the tip of the 

205 Fig. 125. Tracks of a large leopard (male) on wind-hardened and water-logged 

snow; snow has drifted into the tracks. Kedrovaya River in "Kedrovaya Pad' " 
preserve, near Vladivostok. January, 1960. Photograph by A.G. Pankrat'ev. 


Fig. 126. Sketch of foot pads on sole of front (left) and hind paws of a leopard 
killed on Cape Garaov at Vladivostok. 1930, Sketch by G.D. Dul'keit. 

tail is light brown. This background is uniformly speckled with tiny dark 
brown and black spots which do not form rosettes (Fig. 127). With age these 
spots fuse into rosettes. They are larger on the back than on the flanks. The 
underside of the body and the tail are creamy-white and speckled with large 
blackish-brown spots. The weight of a newborn cub is 500 to 700 g and 
body length about 15 cm. The young open their eyes on the 7th to the 9th 
day. On the 12th to the 15th day they begin to crawl around the den and 
by the second month emerge from it to play nearby. At this age they also 
begin to eat meat and shed the juvenile coat. Initially they are fed on milk 
alone, then semidigestexi meat regurgitated by the mother, and finally birds 
and small mammals. After one year the young still differ greatly in 
appearance from adults; in February (1959) the length of the trunk of a 
dead female was 1 17 cm and that of her cub was only 73 cm (Polozov, 1959). 

The cry of a leopard cub is very similar to the meowing of domestic 
cats. Lactation continues for five or six months. As soon as the young begin 
to eat meat the mother takes them with her to hunt. Cubs sometimes 
accompany their mother until she comes into estrus. Young leopards, 
deserted by their mother, continue to live together and follow adult animals, 
consuming the remains of their prey. Hence groups of two to five are 
sometimes sighted. At the end of winter juveniles separate from each other 
and usually abandon the region in which they were bom. The male lives 
with the female only during the period of rut. References in literature about 
the male remaining with his offspring are erroneous. 

Sexual maturity sets in at the age of 2 to 3 years and ability to reproduce 


207 Fig. 127. Cub (male) of an Amur leopard. Sanduga River, west shore of Amur 

Gulf near Vladivostok. February 27, 1956. Photograph by A.G. Pankrat'ev. 

continues up to 12 to 15 years of age. In captivity some animals have 
survived for 21 years. In the London Zoological Garden 52 leopards lived 
on the average of 39.3 months each, but one animal survived for 13 years 
8 months (Mitchell, 1911). 

Molt in leopards has not been studied. 
205 Enemies, diseases, parasites, moriality, and competitors. Leopards have 

no enemies except tiger, and gray and red wolves [dhole]. In India red wolves 
sometimes gather in packs of 30 to attack a leopard, which tries to rescue 
itself by hiding in trees (Anderson, 1964). 

Diseases suffered by wild leopards have not been studied. Only one 
instance of a rabid leopard is known in the Caucasus. In zoological gardens 
this predator suffers from all the diseases characteristic of other feline species 
(infectious enteritis, carnivore distemper, etc.). Three species of helminths 
are known to parasitize leopards in Azerbaidzhan: Taenia hydatigera, 
Mesocestoides lineatus, and Ancyclostoma canium (Asadov, 1947; 
Sadykhov, 1955). These parasites have been reported additionally: Toxocara 
mystax (Mozgovoi, 1953), Dirofdaria granulosa (Skryabin and 
Shikhobalova, 1948), and roundworm Dracunculiis persarum, a 
subcutaneous parasite of man (Pavlovskii, 1935). 

Tiger, gray and red wolves, and possibly striped hyaena, represent serious 
competitors of leopards. In Primor'e it has been noted that at places where 


tigers reside, leopards are absent or very rarely encountered. In India instances 
of leopards being killed by tigers have been recorded. It is known that 
hyaenas chase leopards away from their prey (Pocock, 1939). Oncea leopard 
was mortally wounded by a porcupine (Burl'er, 1955)*. 

Population dynamics. This has not been studied. It is only known that 
due to killing by man and a reduction in its food base (destruction of 
ungulates), the leopard population decreased considerably during the last 
century. In the Caucasus, Tadzhikistan, Uzbekistan, and the Far East this 
species is on the verge of total extinction. 

Field characteristics. The leopard is a large cat weighing 30 to 75 kg. 
It differs from the ounce in its yellow or reddish-yellow background coat 
color, speckled with small, bright, black spots disposed in rosettes over most 
of the body. Its tail is equal to two-thirds of the body length. Its tracks are 
206 smaller than those of a tiger (size of a track on snow 12 cm x 12 cm) but 
far larger and rounder than those of a cheetah. Claw impressions, unlike 
cheetah, are not seen in leopard tracks. Tracks extend in a uniform chain 
with a distance of 40 cm between individual marks (Figs. 121 to 125). 

A leopard conceals the remains of its prey under the roots of fallen trees, 
in patches of wind-felled trees, in upright trees, in hollows, and in rock 
crevices, and feeds on it for two to four days. In Primor'e the head is 
invariably absent from all prey items, even when the carcass has not been 
fully consumed (Vasil'ev et al., 1965). The intestine of victims along with 
their contents are eaten, and even highly decomposed carcasses — which is 
not the case among other species of cats. Leopards bury their feces. 

Leopards are fairly silent animals and their call heard only rarely. During 
the breeding period they call almost every night. At other times their call 
can be heard in the mountains over a distance of several kilometers. The 
leopard's call consists of short, low-pitched "zzugghs" similar to the sound 
of a saw, delivered at short intervals; sometimes the calls are prolonged 
and almost merge with one another. During the breeding period the call 
resembles a harsh cough which tums into labored breathing. On approaching 
a water hole leopards often purr. When disturbed they rumble and snort. 
On falling into a trap the animal maintains silence. (A.S.) 

Practical Significance 

Leopards are fur-bearing animals but their skins are not handled in the fur 
trade of the Soviet Union because the population of this species is too small. 
At the end of the last century in one of the former Krasnovodsk districts 
(western Turkmenia) only 10 leopards were caught annually (Silant'ev, 

*[S'/c]; Bourliere. Not in Literature Cited — Sci. Ed. 


1898). In Turkmenia the following numbers of skins have been prepared: 
4 in 1925-1926, 5 in 1926-1927, 9 in 1927-1928, and 7 in 1928-1929 
(Kogan, 1931). Later the catch in Turkmenia increased: 6 skins were tanned 
in 1948, 7 in 1949, 5 in 1950, 10 in 1951, 6 in 1952, 9 in 1953, 8 in 1954, 
9 in 1955, 41 in 1956, 24 in 1957, 35 in 1958, 19 in 1959.iio In 1912 only 
2 leopards were killed in the Amur region, 11 in Primor'e, 1 in Trans- 
Baikal, and about 15 in northeastern China (Baikov, 1927a). Only 2 or 3 
were caught some 10 to 12 years later in the Ussuri territory (Solov'ev, 1925). 

According to current data (1960's) published by the fur industry the 
skins of this species are not listed separately but grouped with ounce and 
cheetah under the general classification of "leopard". In the 1930's some 
120 "leopard" skins were tanned annually (Kogan, 1931), most of which 
were ounce, and only 20 leopard. The total output of leopard skins per se 
for the Soviet Union up to the 1930's was placed at 20 to 30 pieces per 
annum (Kuznetsov, 1932). In recent decades 20 to 30 skins have been tanned 
in a season but some retained by the people. In the Leningrad fur auction 
of 1956, 40 leopard skins were put up for sale and sold at an average price 
of $10 each (American currency) (Kaplin, 1960). 

The contribution of the U.S.S.R. to the world trade in leopard skins 
is obviously insignificant. The annual world output from 1907 to 1910 was 
10,000 to 1 1,000 skins, of which 5,000 came from Asia and the rest from 
Africa (Brass, 1925). In the 1920's the annual world output dropped to 
between 1,000 and 2,000 (Zhitkov, 1928). In the 1950's leopard skin coats 
207 and other apparel became fashionable and the world output in these years 
soared to 20,000 skins (Shou, 1958). The fur market in Kunming alone 
(Yunnan) handled 4,000 to 5,000 skins annually (1958; A.A. Sludskii). A 
leopard's coak versus one of other skins, reckoned at 100% durability, is 
quite durable with a rating of 75% (opposed to fox with a rating of only 40%). 

Different parts of a leopard's body are widely used in Chinese medicine 
even today and hence, apart from skins, even the bones are marketed in 
China. Limb bones are the most valuable. Desiccated eyes are also highly 
prized. In fact, a hunter often realizes more from the sale of leopard bones 
and eyes than he does from the sale of its skin. 

In the Soviet Union the damage inflicted by leopards on game has been 
very small because the population of this predator has never been large. Only 
recently have leopards been classified as a dangerous enemy of cattle in 
Turkmenia and in the Ussuri territory, but the damage caused by this predator 
is negligible. In the past this cat threatened game even in the Caucasus and 

^"^It is possible tliat some cheetah skins were included in the leopard count 
in Turkmenia. One must also remember that a large number of leopard skins are retained 
by hunters. Up to 15 leopards were killed annually in Kopet-Dag alone in 1940 and 1941 
(V.G. Heptner), 


Trans-Caucasus. Complaints about losses were registered in Lenkoran as 
late as January, 1930 (Starichikhin, 1930). In the Far East leopards were 
formerly responsible for losses among sika deer in state farms. Among the 
enemies of sika husbandry, especially in Pos'et region, leopard ranked 
second, with wolf coming first. Reportedly, a leopard once sneaked into 
a deer park by crawUng through a wire net fence 3.0 meters thick, consisting 
of 16 rows of barbed wire (Menard, 1930). 

In India, where these predators are still quite numerous at places, attacks 
on humans have been reported. In fact some animals have become man- 
eaters to such an extent that they have threatened the safety of human beings 
over a wide area. By the end of 1850's one leopard had killed about 200 
people in a three-year span (Pocock, 1939). In Kumaon in the present century 
two leopard man-eaters killed 525 people before they were put down 
(Corbett, 1957). A leopard becomes a man-eater by accident — when the 
destruction of game by man deprives it of normal food. Moreover, unlike 
tiger, leopards will sometimes eat carcasses. At certain places in India dead 
bodies are abandoned during epidemics; the animal coming across such 
human corpses feeds on them and becomes accustomed to human flesh. 
When the epidemic had ended, it continues to attack living persons. Old and 
208 crippled animals, who cannot hunt ungulates successfully, also turn into 
man-eaters. Man-eaters have also been reported in China but are extremely 
rare (Allen, 1938). 

In the Soviet Union neither biological nor social conditions have given 
rise to man-eaters either in the past or present. Usually only an injured leopard 
will attack man (very few cases); nevertheless instances of an uninjured 
normal animal attacking man have been recorded. In the Caucasus during 
the winter of 1881/1882 in the Groznyi region a leopard tore a man to bits 
in the forest. In 1875 near Sochi one attacked a sleeping worker, severely 
bit his occiput, and attempted to carry him away into the forest, but 
fortunately was repulsed by other workers (Radde, 1899). In the 1890's 
in the Lenkoran lowland near Prishib village another leopard killed a 
15-year-old shepherd boy (Dinnik, 1914). 

In Turkmenia two incidents have taken place. Early in the present century 
in a gorge in western Kopet-Dag, a leopard unexpectedly ambushed a man 
from behind a clump of rushes. In the life-and-death struggle which ensued 
the man managed to strangle and stab his attacker. It was later discovered 
that the leopard was old with worn-out canines. The hunter was further saved 
from serioiLS injuries partly because he had on a heavy coat [khalat]. In the 
early 1920's a hunter from Verkhne-Skobelev village in hot pursuit of rock 
partridge in the Kopet-Dag jumped onto a stone behind which a large leopard 
happened to be sleeping. The animal sprang instantly to the attack and, 
placing its forepaws on the hunter's shoulders, tried to seize him by the 


throat. Luckily, the gun with the hammer cocked, which the hunter was 
carrying in his hands, jammed in the attacker's throat, and the predator died 
within a few seconds. In my opinion (V.G. Heplner) this incident should 
not be viewed as a leopard attack. By and large, leopards are far less 
dangerous to man in the Soviet Union than wolf; this predator has killed 
quite a number of people in this country (see Volume II, Part 1 of the present 
monograph. In the Kopet-Dag, a leopard has sometimes tracked a man as 
though pursuing him, but has not attacked (V.G. Heptner). 

In the Caucasus leopards were shot on chance encounters or caught 
in traps set up mainly in trees standing in water or on specially made bridges. 
Strychnine administered in the form of pellets embedded in the remains of 
prey was also used. In Turkmenia leopards were shot and caught in traps 

208 Fig. 128. Catching of live leopards in the Far East. Hunter nudging one taking 

refuge in a tree with a pole (from Sysoev, 1955). 


set up on trails. In the Far East native people were as afraid of leopard 
as of tiger. Only rarely was one caught by setting up bows and guns on 
trails and at the site of remains of a kill. The animal was also shot in chance 
encounters, caught in traps, poisoned with strychnine, and chased on fresh 
snow with the help of dogs. In the 1920's some deer farmers used German 
209 sheepdogs for hunting leopards in deer parks and near them. These dogs 
wore spiked neck collars to protect them from seizure by the predator. Two 
or three dogs spotted an animal and chased it up a tree where it was then 
shot. In the hunting season of 1927 one hunter (A.A. Bogoyavlenskii) with 
two dogs caught three leopards (V.V., ,1927; Vereshchagin, 1929). A leopard 
quickly kills pursuing dogs by biting their skulls. The predator is highly 
resistant to wounds and it is a rare feat indeed when one is felled with a 
single shot. 

Fig. 129. Leopard nudged down from a tree and pinned to the ground by hunters 
(from Sysoev, 1955). 


In the All-Union standards on skins in the 1940's to the 1960's leopard 
skins were classified according to size, grade, and flaws. The raw skin of 
a leopard in the last decade was not prized and, depending on grade, sold 
for 80 kopecks to 4 rubles 70 kopecks. Formerly, it was more expensive; 
for example, in the 1860's in the Ussuri territory a good skin sold for 15 
to 20 silver rubles (Przheval'skii, 1870). Skins were used in making rugs 
and sleigh robes. Apart from skins, in the Far East the Chinese formeriy 
valued other body components (meat, bones, blood, brain, bile, teeth, and 
claws). A male was priced higher than a female and an adult higher than 
a cub. The average price of a freshly frozen carcass was two to three gold 
rubles per kilogram. Different parts of the carcass were used in Chinese 
medicine and as talismans. 

Leopards, like tigers, are especially honored by natives of the Far East 
and always worshipped. Spotted idols used to be set up on altars in high 
altitude passes (Baikov, 1927). Udekheits and Orokhets of olden times 
idolized this predator and its idols were found among them (Schrenk, 1858; 
Przheval'skii, 1870). 

In recent decades, due to the great demand for live leopards, they have 
been trapped or caught live without traps. In the Ussuri territory hunters 
catching tigers (see section 'Tiger") also caught leopards. On finding fresh 
tracks of a leopard, hunters set their dogs in pursuit. When the dogs have 
stopped the animal or chased it into a hiding place, hunters pin it to the 
ground with forked poles and then truss it (Sysoev, 1955). If a leopard escapes 
the dogs and climbs onto a tree, the hunter nudges it down with a pole and 
his companions pin it to the ground. Not only young leopards but also adults 
are caught with forked poles. Leopards survive and breed well in captivity. 

In the Soviet Union this big and beautiful predator, like the tiger, is 

an asset to the mountains and forests in which it lives. At present it causes 

no harm to cattle or game. It also poses no danger for man if not attacked. 

no Leopards should be protected in the Soviet Union since this species is of 

great scientific and aesthetic value. 

To protect the leopard and prevent its total extinction, hunting and 
catching live animals should be banned throughout the Soviet Union.^^^ 
International conventions for the conser\'ation of leopards should be 
established with neighboring govemments (Korean Democratic Republic, 
China, Afghanistan, Iran, and Turkey). Reserves and preserves ought to be 

'^'Such a measure had already partly arrived in Primor'e territory in 1956 and shooting 
of leopards banned year-round in Khasansk, Vladivostok, Shkotov, Budenov, and Lazov 
regions, and within Vladivostok, Artem, Suchansk, and Nakhodka municipalities. A penalty 
of 100 rubles is now imposed for catching a leopard. Contrarily, in other republics (Armenia, 
Azerbaidzhan, Turkmenia, and Tadzhikistan) killing leopards continues year-round and a 
prize is even awarded for any animal killed. 


set up for the conservation not only of this predator, but also of the animals 
on which it survives (mountain sheep, roe deer, sika deer, etc.). Raising 
leopards should be organized in zoological gardens by creating pedigree 
groups of a single subspecies. 

In Africa this predator has almost totally disappeared over large expanses 
in recent years because leopard skin coats became fashionable in the 1950's 
and 1960's. Subsequently wild pig and baboon, the main food items of 
leopards, multiplied unchecked; the loss inflicted on crops by these animals 
far exceeded any loss to domestic animals caused by leopards in the past. 
For this reason, and under intensive pressure from campaigning ecologists 
in several African states, the leopard was removed from the category of 
dangerous animals and placed on the list of animals to be protected by law 
(Arrua, 1958*). In some countries (Kenya) work is now in progress to 
reacclimatize the leopard (Right, I960**). (A.S.) 

'[Sic]: not in Literature Cited — Sci. Ed. 

'*[Sic]; Wright? Not in Literature Cited— Sci. Ed. 

Genus of Snow Leopard, or Ounce [Irbis] 
Genus Uncia Gray, 1854 

21 1 1854. Uncia. Gray. Afin. Mag. Nat. Hist., 14, p. 394. Felis uncia Schreber, 
Large-sized animals. 

Legs not long, trunk very long, general build stocky, sacral region high, 
higher than shoulders, and profile of back slopes forward somewhat 
(appearance typical of small cats, Felis; see Fig. 30). Tail length distinctly 
more than half length of body. Claws wholly retractile and paw structure 
same as in big cats (Panthera, see Fig. 278). Body covered with dark annular 

Skull quite powerful with tuberosities, crests, and powerful zygomatic 
arches, but less massive, less heavy, and with less developed crests than 
genus Panthera. Skull broad and short with large orbits set high; interorbital 
region distinctly elevated, with upper line of profile sharply curved, falling 
relatively steeply fore and aft of interorbital region. In general, skull bulges 
notably. Facial region shorter than in big cats (genus Panthera); distinct 
notch occurs at places where nasals join frontals. Nasals short and broad; 
their anterior width almost equal to their length along midline. Basicranial 
axis markedly raised toward the front (not horizontal), meets basifacial axis 
(axis of facial portion of skull) and forms a distinct angle. Postorbital 
constriction distinct and broad (broader than interorbital). Cranium relatively 
more voluminous and enlarged than in big cats (genus Panthera), not 
elongated, or slightly so, with facial region of skull relatively less powerful. 
Orbits open. By and large skull resembles that of small cats (genus Felis). 

Tympanic bulla relatively small and slightly expanded. Distance between 
it and posterior wall of glenoid fossa small; the two almost touch each other. 
Antero-outer (ectotympanic) chamber of bulla very well developed; two 
chambers nearly identical in size, with suture between them shifted away 
from auditory meatus (Fig. 23). Anterior part of entotympanic chamber very 
narrow in front. Deep depressions present on basioccipital portion of occipital 
bone in front of jugular foramen on each side, internal to tympanic bulla. 
Structure of hyoid apparatus same as in big cats (genus Panthera), i.e., the 
epihyal in the form of an elastic ligament (Fig. 31). Chin portion of lower 
jaw set steeply (almost at a right angle). 

Dentition complete (second premolar present); teeth, especially canines, 
relatively less massive than in big cats (leopard). 


Sexual dimorphism absent. 

This predator generally catches prey as large or even much larger than 
itself. It hunts by lying in wait. It is a solitary animal and lives in mountains, 
212 mainly in high altitudes up to snowline. In spite of the structure of the hyoid 
apparatus, similar to that of big cats (genus Panthera), the "roar-growl" 
type of call is not heard. It "purrs" during both inhalation and exhalation 
(as in small cats, genus Felis). It tears its prey in the manner of big cats 
but its eating posture is that of small cats (genus Fells, see Fig. 32). 

The range of the genus (see description of the species) is relatively 
small, somewhat less than that of other genera. It covers montane Central 
Asia as far as the Himalayas in the south, in the west to the Hindu Kush, 
Pamir-Gissar system, and Tien Shan, in the north to the Altai (probably 
to [lake] Baikal in the past), and in the east includes eastem Tibet. 

The genus Uncla is unique in several respects and placing the ounce 
in genus Panthera is not justified. Craniologically it differs from species 
of the genus Panthera more than the latter differ among themselves in any 
combination. At the same time, genus Uficla shows a combination of several 
moфhological and ethological characteristics of both big cats (genus 
Panthera) and small cats (genus Fells). Thus the structure of the cranial 
region of the skull is extremely similar to that of large species of the genus 
Fells, but a hyoid typical of genus Panthera, etc. Ethologically, as far as 
can be judged from observations of captive animals, the ounce is closer 
to small cats {g&xms Fells) and essentially identical with their "peripheral" 
species — the clouded leopard (F. nebulosa; Hemmer, 1964 and 1966). This 
species has also been placed time and again in "subfamily Pantherinae," 
but it is now clear that this is a genuine small cat although it shows some 
specialized features. It is sometimes placed in a separate genus (Neofells) 
even by authors who recognize only a few genera in the family (Ellerman 
and Morrison-Scott, 1951 and 1966). Features of the ounce, discussed 
above as an intermediate form between genera Panthera and Fells s. 1., 
define its taxonomic position and, simultaneously, highlight the difficul- 
ties of a clear separation of genera of present-day cats (excluding the 

Paleontological data pertaining to the origin of the genus are not avai- 
lable. Quite possibly its origin, like that of the genus Panthera, should be 
placed at the end of the Pliocene or in the Pleistocene. 

There is only one species in the genus, ounce or snow leopard, Uncla 
uncla Schreber, 1775, representing 3.5% of the species of the family. 

The ounce is a valuable game and fur-bearing species and one of the 
prized possessions of zoological gardens. It exerts a control over the 
population of some mountain ungulates. 

In the USSR the one species in the genus constitutes 9.0% of all species 


of the family inhabiting the Soviet Union and 0.3% of mammals of our 

The geographic range in the Soviet Union covers the montane regions 
of the eastern parts of Middle Asia and part of the extreme south of middle 

This species is an object of sport and also important to the fur industry. 
It is mainly valued, however, as an object of trade in live animals for 
zoological gardens. The damage inflicted by the ounce on livestock is very 
little and can be ignored. (V.H.) 


Uncia uncia Schreber, 1775 

1775. Fells uncia Schreber. Sdugeth., tab. 100, vol. 3, p. 586 (1778). Type 

locality not indicated. 2 
213 1830. Fells Irbls. Ehrenberg. Ann. Sc. Nat., vol. 21, pp. 394 and 406. 

Substitute for F. uncia Schreber. Altai. 
1855. Fells uncloldes. Horsueld. Ann. Mag. Nat. Hist., vol. 16, p. 105. Nepal. 
1950. Panthera uncia schnelderl. Zukowsky. Zool. Gart., vol. 17, p. 213. 

Sikkim. (V.H.) 


Uficla uncia is the only species of the genus. 


Size large, roughly corresponds to that of leopard (Panthera pardus), but 
slightly smaller on the average. 

Trunk very long and stocky; general appearance similar to that of small 
cats (genus Fells, Fig. 30), i.e., with a slightly raised sacral region. While 
generally resembling the leopard, the ounce also differs in its less strongly 

^For a correct Russian name for this species, see footnote no. 1 under "leopard". From 
a scientific viewpoint it is incorrect to call this animal simply "leopard" [bars]. 

"S.I. Ognev (1935; followed by Stroganov, 1962), based on some indirect evidence, 
suggested that Iran should be regarded as the type locality. His conclusions are not adequately 
substantiated; in particular the occurrence of ounce in Iran has not been generally proved; 
evidently this animal is absent there or is only found as an exceptional rarity (vagrant — ?, 
see "Geographic Distribution"). In any case, Iran as the tjpe locality is inadmissible. The 
far better substantiated suggestion of R. Pocock (1930), to consider the Altai the type locality 
not only for the Soviet Union but also Mongolia [sic], should be accepted. (V.H.) 


developed anterior (chest) portion of the body and a relatively small and 
more rounded head. Upper line of profile of head bulges, eyes quite large, 
and pupils round. Ears short with rounded tips, broad at base, almost hidden 
within the coat in winter; long hair (tuft) at tips of ears absent. Mane or 
long hair on cheeks (whiskers) also absent (Fig. 130). Vibrissae white and 
black and up to 105 mm long. Tail very long, relatively longer than that 
of other Soviet cats, and more than three-fourths length of body. Paws broad 
and massive; claws a light horn color. In spite of having dense and long 
hair, the animal with its somewhat flattened and less muscular trunk appears 
less massive than the leopard (Figs. 132 and 133). Forepaws larger and 
more powerful. 

The winter coat is very rich, soft, and silky. Tail covered with very 
long and dense, lax hairs, and appears very thick and fluffy. 

Pelage of the ounce, compared to that of other species, moderately dense, 
long, and soft. On the back, density about 4,000 hairs per cm^, with eight 
underfur hair to one body hair. A characteristic feature is the absence of 
any significant difference in the length of different categories of hair; guard 
hair only 1.2 times and body hair of category I just 1.1 times as long as 
hair of underfur. Body hair of category IV and especially that of category 
III somewhat longer than body hair of categories I and II. Length and 
thickness of guard hair on the back 54 mm and 77 microns respectively; 
length of body hair (categories I to IV) 48, 47, 51, and 49 mm and thickness 
76, 58, 43, and 34 microns; length of underfur hair 43 mm and thickness 
20 microns. Type of fur abdominal"* (B.F. Tserevitinov). 

In pelage density the ounce differs from all other big cats and is more 
similar to small cats (genus Felis), occupying a somewhat intermediate 
position between them. Its being "intermediate" is important in any 
comparison with small cats. These differences, however, are hardly important 
in taxonomy and are more associated with conditions of the habitat of the 
animal and its thermal requirements (thermoregulation). The contrast between 
winter and summer coats is very great, far more than the corresponding 
contrast in the leopard. 

General color of the main background of winter coat very light gray, 
almost white, or whitish with a smoky tinge, more perceptible on back and 
up the flanks; it may also be light yellowish. Lower part of flanks, whole 
of underside of body, and inner surface of legs even lighter, being almost 
white or with a yellowish tinge (Fig. 132).^ 

*Initial quotation mark for this quote missing in original Russian; meaning of last 
sentence unclear. — Sci. Ed. 

^he yellowish shade seen in skins of some animals which died in captivity 
(Zoological Museum, Moscow University) is possibly an artifact. 




215 Fig. 131. Skull of snow leopard, Uncia uiicia Schreb., No. S 85724. Collection 

of the Zoological Museum, Moscow University. Trans-Ili Alatau, Tien Shan. August 
4, 1969. Sketch by N.N. Kondakov. 

There are no distinctly yellow or reddish or tan shades in the color 

215 scheme of the ounce. On the main background are scattered dark spots, 

some of which are annular, with a diameter of up to 7.0 to 8.0 cm, and 

others large or small and solid (Fig. 133). Spots are sparse, far more sparse 

than in the leopard. 



Fig. 132. Snow leopard. Vil'nyus Zoological Garden. Photograph by B.B. Marm. 

Solid spots of different sizes are scattered on the head (smallest) and 
neck and legs (larger ones above, smallest below) where annular (rosette) 
spots are absent. The latter are rounded or somewhat elongate and disposed 
on the back, flanks, upper portion of the thighs, along the shoulders, and 
on top of the proximal portion of the tail (sometimes in two rows here). 
On the posterior part of the back, spots often fuse into short transverse stripes. 
Between the annular spots a few small solid spots are scattered. Large solid 
spots on the distal half of the tail often surround it in the form of an 
incomplete ring set in a transverse direction; the very tip of the tail is usually 

The color of the central areas of annular spots corresponds to the main 
background in a given section of the body. The spots are black but their 
outlines are not sharp in a dense coat, and are often greatly blurred; their 
color is therefore not complete and appears more as a dark gray shading. 
This is particularly true of the annular spots. Solid spots, especially the 
large ones, are more shaфly outlined and a more intense black color. In the 
216 summer coat the general background appears more light-colored, almost 
white, and the outlines of the dark spots sharper, darker, and usually pure 



Fig. 133. Snow leopard. Vil'nyus Zoological Garden. Photograph by B.B. Marm. 

black. The smoky film is fainter than in the winter coat."* 

The pattern of spotting shows some individual variability in density of 
spots, etc. The color of the spots may also vary (Figs. 134 and 135). Usually 
they are pure black but sometimes brownish. The main background color 
of the coat is less variable. In general, however, individual color variability 
in both these features (color and pattern) is comparatively less marked in 
the ounce than in the leopard. Sexual dimorphism in color is absent. 

The young in their first coat (newbom) differ sharply in the depth of 
pigmentation of spots, which are few in number, and also in the presence 
of few annular spots. Large black or brownish solid spots are present on the 

'*The characteristics of the summer pelage of the ounce and its variability is not 
adequately known. The animal carries it for a very short time and most pelts in museums 
are of winter skins. Usually, therefore, winter skins are described. 


218 Fig. 134. Variation in color in winter coat of snow leopard, Uncia uncia Schreb. 

1— No. S 35130, female, February 1, 1939. Dzhety-oguz, 55 km south of 
Pi-zheval'sk, Tersk-Alatau, western Tien Shan; 2 — No. S35128, male (not adult), 
February 1, 1939, same places as above; — No. S 35126, female, February 1, 1939, 
Dzhetyoguz; 4 — No. S 72736, male, December 19, 1964, western Tien Shan; 
5 — No. S 72958, female, March 20, 1965, western Tien Shan; 6— No. S 35127, 
female, February 1 , 1939, Akshiprak area, 90 km southwest of Przheval 'sk, Tersk- 
Alatau, western Tien Shan. All skins from collection of the Zoological Museum, 
Moscow University. Photograph by V.G. Heptner. 

219 Fig. 135. Variation in color in summer coat of snow leopard, Uncia iincia Sclireb. 

1 — No. 49243, August 13, 1939, Tien Shan; 2— No. 49240, female, June 13, 

1940; Tien Shan; 3 — No. 49953, female, 1945, Mongolia; 4— No. 49242, male, 

August 8, 1939, Tien Shan. All skins from collection of the Zoological Museum, 

Moscow University. Photograph by V.G. Heptner. 


back and similar short, longitudinal stripes in the rear (see Fig. 150). 
Differences from the leopard are quite distinct. By the first winter the coat 
of cubs does not differ from that of adults.-'' 

Geographic variation in color is absent. 

Apart from the features discussed above (see characteristics of the 
genus), the skull of the ounce differs from that of leopard, with which it 
bears features of similarity, in the following respects. Skull relatively broad 
and short with widely separated zygomatic arches; cranium more enlarged 
and roomy with a relatively broad postorbital region (its width about 30% 
of condylobasal length and usually more than interorbital width), reflecting 
affinity with small cats (genus Felis) and not big ones (genus Panthera); 
and skull wider in interorbital region. The above results in a more rounded 
skull which, on the whole, is similar to that of small cats. Orbits placed 
somewhat higher, and more rounded; zygomatic process of squamosal stops 
far short of base of postorbital process of jugal; lower anterior edge of orbits 
217 not thickened; hard palate shorter and broader; rostrum broader but its width 
above canines not greater than postorbital width; and choanal edge of 
palatines with two small pointed projections along sides of suture between 
palatines. Anteriorly, nasal process more rounded; interpterygoid vacuity 
wider; paroccipital processes thin and long, project markedly beyond surface 
of tympanic bullae, and tips pointed and bent forward in form of a hook. 
Jugular foramen relatively small and rounded. 

Measurements for adult animals (males and females combined): body 
length (10) 103 to 125 cm (M 112.1); tail length (10) 80 to 105 cm (M 
91.8); length of hind foot (8) 22.5 to 26.0 cm (M 23.8); and length of ears 
(8) 6.6 to 7.2 cm (M 7.0). Weight (5) 22.0 to 39.0 kg (M 33.0) (material 
from the Zoological Museum, Moscow University, and data of A. A. Sludskii 
and B.B. Marm). 

Female (Talassk Alatau, April): body length 125 cm, tail length 90 
cm, hind foot length 24.5 cm, ear length 6.6 cm, height at withers 50 cm, 
height at sacrum 60 cm, and weight 39 kg. 

Maximum body length up to 130 cm and weight up to 40 kg (Stroganov, 
1962). Females are evidently somewhat smaller than males. 

Cubs aged about 9 to 10 months (January and February): body length 
85, 82, 87 cm; tail length 75, 65, 76 cm; length of hind foot 22, 21, 22 
cm; and ear length 68, 63, 70 cm (first and last from the same litter; 
Zoological Museum, Moscow University). 

Skull measurements are given in Table 4. (V.H.) 

^The view that the spots of ounce become more diffuse with age (Stroganov, 1962) 
needs confirmation. It may be true of the first adult coat but is hardly applicable to subsequent 



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Systematic Position 

Uncia uncia is the only species in the genus. 

Geographic Distribution 

This species is distributed in the upper and middle zones of mountains in 
Central Asia, Turkestan, and southern Siberia. 

Geographic Range in the Soviet Union 

The range in the Soviet Union (reconstructed) represents the northwestern 
and northern periphery of the range of the species and covers the montane 
systems in eastern Middle Asia (Turkestan) and southern Siberia from the 
Amu-Darya to Baikal. 

In the west the range of the ounce covers the Pamir-Gissar system and 
Tien Shan, the entire Pamir, the Darvaz range including its southwestern 
spurs, the Peter the Great, Trans -Alaisk, and Gissar ranges together with 
the Baisuntau mountains, the Zeravshan range westward to the districts of 
220 Pendzhikent and Turkestansk to the west, at least to the meridian of Zaamin 
(Zaamin preserve) and the Alai range. Ounces were never present in the 
Mal'guzar mountains (Fig. 136). 

The southern boundary fronting on the Amu-Darya in southern 
Tadzhikistan describes an arc beginning at Pyandzh on the north and 
encompassing the Kulyabsk, Dashti-Dzhumsk (connecting the Khazrat- 
Ishi mountains with the Darvaz range), Muminabadsk and Kyzyl-Mazan>k 
regions, where the animals are regularly met with. From here the boundary 
runs northwest and encircles Dushanbe from the north. North of that city, 
the ounce lived along the Varzob. From there the boundary runs along the 
southern slope of the Gissar range to the west and later southwest along 
the Baisuntau [range]. In Baisunsk mountains the ounce is distributed in 
the south at least to the latitude of Deinau on the Surkhan-Darya. This species 
is evidently absent at Kugitang. In this part of its range (southern 
Tadzhikistan), the animal occupies the higher portions of the country. It 
vicariates with the leopard, which occurs here in several places, but to the 
south of the range boundary described above, and at lower elevations. 

Farther to the north and northeast, the ounce is met with in all the ranges 
of the Tien Shan system, to the south including the Kuraminsk and Ferghansk 
ranges, reaching the Ferghana valley from the north, and westward reaching 
the western spurs of the Chatkal'sk (Montane-forest preserve near Parkent), 
Pskemsk, Ugamsk, and Talassk ranges (in the west to Aksu-Dzhebagly 
preserve). The Karatau, Kirgizsk (Aleksandrovsk), Kungei Aatau and Trans- 


Hi Alatau ranges form the northern limit of occurrence (ounces are 
encountered on the northern slope south of Alma-Ata). In the Chu-Ili 
mountains ounces are absent; they inhabit the Dzhungarsk Alatau and its 
offshoots — the Altynemel', Katutau and Aktau — and the Tarbatai and Saur. 

In Altai, the ounce occurs far to the south. In the west they have been 
met with along the upper and middle course of the Bukhtarma from its 
uppermost reaches (Beleya ВегеГ River and Chindagatui mine) and noticed 
even as far as the lower Zyryanovsk. They also live on the upper Narym, 
between Narym and Bukhtarma, and in the Sarymsakty range south of 
Katon-Katagai (Altaisk station). The more easterly range includes the 
mountains of the Argut basin (upper Katun), including its tributaries the 
Dzhassater in the south and Shavla to the north (near the mouth of the Argut 
on the Katun). The Chuisk steppe and the surrounding ranges, and evident- 
ly the mountains of the entire Chui basin also fall within the range. Even 
more eastward the range covers the Bashkaus basin, the left tributary of 
the Chulyshman, and Chulyshmansk basin in the north, including the 
Chul'chu and Kygu, its right tributaries. Here (Altai preserve) the range 
extends to the southern extremity of Lake Telets. Thus, the following main 
ranges of the southern and, in part central, eastern, and northeastern Altai, 
and associated massifs fall within the geographic range: Narym, southern 
Altai, Tabyn-Bogdo-Ola, Listvyaga, Katunsk, Sailyugon, southern and 
northern Chui; Chikhachev, Aigulansk, and Chulyshmansk mountains; 
Chulyshmansk uplands; and Shapshal'sk mountain (Chapchal). 

In the portion of Altai outlined above the distribution of the ounce 
is uneven — in the far south they are found more or less regularly (upper 
Bukhtaria, Chuisk mountains), and in the north and northeast, occur as 
vagrants, very rarely at some places (Lake Telets). 

Still farther east closely associated with the Altai-Sayan system and Tuva 
Autonomous District, the distribution includes the Tannu 01 range, region 
of the upper Kemchik, and the upper Bol'shoi and Malyi Yenisei. Farther 
north, in the Sayan range, directly adjoining the Shapshal'sk [range] (Altai), 
the distribution runs to the east at least to the upper Kazyrsuk (right tributary 
of the Yenisei'^). It is highly probable that the range included the entire 
western Sayan system (except the Sayansk, Kurtushibinsk, and Ergak- 
Torgak-Taiga ranges), and the eastern part of eastern Sayan system — Udinsk 
range (source region of the Bol'shoi Yenisei). 

Farther east, the range covers Tunkinsk [range] and possibly the Юtoisk 
bald (west of the southem end of Baikal) and also Pri-Baikal mountains along 

^One was caught approximately 130 km south of the city of Abakan, 40 km from Beya 
(Beiskoe) village. References to occuirence in the first half of the last centurj' at Krasnoyarsk 
(Radde, 1862) are dubious. Probably the region in a broad sense is intended. 


223 the Angara and upper Lena in the north at least to Balagansk.'^ This is the 
northernmost point of the eastern part of the range. 

Toward the middle of the present century and moreover, even in the 
last century, the range of snow leopards shrank at some places. Thus ounces 
were already absent in Karatau or probably only wandered into the southern 
part of the range; they were absent in the Sayans (though likely intniders 
from the Altai into the westernmost parts), on the Tunkinsk balds (still 
possibly vagrant from Mongolia), and in Pri-Baikal. Ounces occurred in Pri- 
Baikal more as vagrants even in the eighteenth century. In the rest of the 
range the outlines given above are still maintained, although some peripheral 
portions are only sites of fairly regular intrusions.^ 

Some false or extremely dubious views crept into literature about the 
distribution of the ounce, and were long maintained and sometimes still 
supported. The report of Pallas (1811) about the occurrence of the ounce 
along the Olekma, a tributary of the Lena, 200 versts from its mouth, is 
based on a very rare intrusion, or possibly is even a mistake (false 
understanding concerning tiger). Pallas obtained this information from the 
diary or manascript of the elder Gmelin ("as I found in the rough notebooks 
(in adversariis) of the elder Gmelin"). 

The statement of Pallas (1811) that the ounce probably occurred along the 
Uda River and between it and the Amur River had very large consequences. ^ 

^Reference from Pallas (1811). This information is so positive that it is hardly possible 
to contradict the occurrence of the ounce in Pri-Baikal (Ognev, 1935). Mountain sheep 
evidently lived foiTnerly in Pri-Baikal and the presence there of Siberian ibex was also possible. 
The range of the ounce in Siberia mostly coincides with the range of these species (see Volume 
I of the present monograph). 

^Range according to data of Pallas, 1811; N.A. Severtsov, 1873; S.A. Seveitsov, 1929; 
Selevin, 1929; Ognev, 1935; V. Skalon, 1936; Yurgenson, 1938; Levnev, 1939; Kolosov, 
1939; ShuPpin, 1948; Chernyshev, 1950; Yanushevich, 1952; Sludskii, 1953; Ishunin, 1961; 
Stroganov, 1962; and others; and original material of V.G. Heptner. 

In outlining the distribution in Tuva and other regions farther east, the data of Pallas 
(1811) have been specifically used. According to Pallas (original in Latin), the ounce (which 
he called Felis pardus) "... intiuded from Central Asia along the montane forest deserts 
at places in .southern Siberia here and there [passim], was seen more rarely in the Altai 
mountains, and more often at the sources of Yenisei proper and Kemchik (Kemchug) and 
in the mountains around Baikal ... On the Lena it was often caught in the vicinity of the 
city of Balagansk on the Lena ... A stuffed animal is preserved in the Academy Museum; 
the skin is that of an animal killed around Tunkinsk at Baikal" (pp. 17 and 18). This 
information has been ignored or contradicted by some authors without adequate justification, 
mainly due to ignorance of the original text. 

^According to the original text of Pallas (1811, pp. 17 and 18, translated from the 
Latin), "They say the animal was reportedly encountered (dicitur frequens in the region 
of the Uda (Uth) River and between it and the Amur. Yakutians going there from the Lena 
were afraid of it and called it tenible (terribilem) to the utmost extent (ka-chai) and on 
encountering its tracks did not dare cross them." 


Some earlier workers investigating the Amur and Ussuri regions (Mid- 
dendorff, 1857; Maak, 1859; Schrenk, 1859; Radde, 1862; Przheval'skii, 
1870; A. Nikol'skii, 1889), evidently without skins or skulls in hand, 
presented all their information on leopard as pertaining to ounce, relying 
entirely on the authority of Pallas. It should be remembered that Pallas merely 
passed on this information with extreme caution, prefacing it with * 'They 
say. ' ' Even after him, as quite often happens in copying from another author, 
the assumption turned into an affirmation (Zhitkov, 1936) and has been 
used as such to our time. Even some specific references to the Stanovoi and 
Yablonovyi ranges, which Pallas never mentioned, began to appear in the 
literature. From the account given above, it is quite clear that Pallas reported 
the mistaken accounts pertaining to tiger, ^° such as the instance of an 
intrusion from the Olekma reported from Gmelin. Tigers in more recent 
periods have moved still farther north. 

The situation is similar in the extreme west of the range under 
consideration. Information about the occurrence of the ounce in the Kopet- 
224 Dag, apparently quite convincing (Khristof, 1882"'; Zarudnyi, 1891*; 
Satunin, 1905; Bil'kevich, 1918; Ognev, 1935; and some others), and quoted 
extensively even in textbooks, is incorrect. A special study conducted in 
the Kopet-Dag (V.G. Heptner) revealed that the ounce is absent there. All 
affirmations are evidently based on dubious answers to queries in a study 
of the particularly rich and very light-colored winter skins of leopards (P. 
pardiis). The ounce is, of course, absent on the Serakhsa (Zarudnyi, 1891*) 
and conditions there are not favorable for its residence. The mounted snow 
leopard exhibited in the Ashkhabad museum was prepared from a skin 
brought by S.I. Bil'kevich from "Eastem Bukhara" (V.G. Heptner). After 
almost 90 years of studies in Turkmenia, not one skin or skull of snow 
leopard has been found. 

To resolve the question of the ounce in Turkmenia, it is important to 
note that the question of its occurrence in Asia Minor and Iran, especially 
in districts adjoining Turkmenia, as shown by special studies (Pocock, 1930 
and 1939), have yielded negative results. The latest information for Iran 
(Misonne, 1959; Lay, 1967) is also so confused and indeterminate that 
zoologists have acknowledged the absence of proof for its occurrence in 
that country. In foreign literature about Iran reference is sometimes made 
to the data of Ognev (1935) about the occurrence of this animal in the 
Russian part of the Kopet-Dag as conclusive, and based on this the possibility 
of its existence in the Iranian Kopet-Dag assumed. It should be kept in mind 

^•^Middendorff (1857, p. 75) similarly comments that in the case of the Tyrma, the 
account pertained positively to tigers. Middendorff himself found no signs of the occurrence 
of the ounce in the Stanovoi range. 

•■'Not in Literature Cited — Sci. Ed. 


that leopards in southern Iran are particularly light -colored, almost white, 
especially in winter, which may have confused contemporary and past 
(Blanford, 1876) authors. It is significant that in Afghanistan the ounce 
inhabits only the extreme westem, much higher parts of the Hindu Kush 
adjoining the Pamir (Badakhshan and Bakhan; Kullmann, 1965, 1967, and 

Geographic Range Outside the Soviet Union 

The range outside the Soviet Union (Fig. 137) covers the Mongolian and 
Gobi Altai, Khangai and the montane district of Lake Kosogol [Hobsogol] 
in the Mongolian Peoples Republic, Tibet to the Altyntagh on the north and 
Kam in the east, and the Himalayas from the eastern end to Kashmir. In 
225 Afghanistan this species inhabits the Hindu Kush. Here it occupies only 
the extreme eastern portions (see above). The leopard inhabits much lower 
levels there, mainly in the west and north, and relations between the two 
species are the same as in southern Tadzhikistan. (V.H.) 

224 Fig. 137. Reconstructed species range of snow leopard, Uncia uncia Shreb. 

V.G. Heptner. 


Geographic Variation 

Ounces do not exhibit geographic variation, or else it is very insignificant. 
A distinct form, U. u. imcioides Horsf., 1855, has been described from Nepal, 
but this separation was done without adequate substantiation. Nevertheless 
some contemporary scientists favor recognition of the Nepalese form 
(Haltenorth and Trenze, 1956). On the other hand it has been shown (Pocock, 
1930 and 1941) that Himalayan animals do not differ from those of the Altai. 
Apparently this is correct. The feature suggested as a diagnostic character 
(color) falls in the category of individual variation. The same applies to the 
form schneideri Zuk., described from a photograph of a skin (!). 

The absence of clear geographic variation in the ounce is due to the 
relatively small range of the species. It is an extremely stenotopic species 
and everywhere in its range is restricted to identical conditions of existence. 
If in spite of this a taxonomic separation of Himalayan animals is proved, 
the name U. u. iincia Schreb., 1775 should be assigned to the Russian form. 


Population. The ounce is rare in the eastern Pamir (Tadzhikistan) but 
common in the western sector. They are fairiy common in the Alaisk, Gissar, 
Zeravshan, Turkestan, and other ranges. In recent years in Tadzhikistan 
about 60 are trapped and caught live each year. In Uzbekistan this predator 
is not rare in the Ugamsk, Pskemsk, Chatkal'sk, Gissar, and Turkestan 
ranges, but is definitely rare in the western and southern spurs of the Tien 
Shan. The ounce is common in most of the Tien Shan ranges in Kirgizia. 
About 30 to 40 animals are captured there every year. In Kazakhstan this 
predator is extremely rare in the Karatau, but common in the Talassk Alatau 
and other ranges of the Tien Shan. Here about 10 to 15 animals are now 
caught annually. The animal is rare in the Tarbagarai and extremely rare 
in the Saur, southern (Kazakhstan) and central Altai, and also in the western 
Sayan and Tannu-Ola. In the latter, only solitary animals are caught and 
these not every year. 

Habitat. In summer the ounce is confined to higher levels in the majority 
of mountains, mostly in the subalpine and alpine zones, and often is 
encountered at the limit of permanent snow. In the high Alichur, in the Pamir, 
tracks are sometimes detected, even in winter, at a height of 4,500 to 5,000 m 
above sea level (R.N. Meklenburtsev). Ounces at that time are confined 
to mountain crests where the snow has been blown off by the wind. In the 
Gissar range, snow leopards live in the high mountains where rocks abound, 
but descend in winter into the shrub junipers along with the ibex, but not 


below this level (Leviev, 1939). On the northern slopes of the Turkestan 
range ounces have been observed in summer only in the upper half of the 
juniper belt, commencing roughly at a height of 2,600 m or more above 
sea level. Here they are confined to rocky sites and do not leave the 
shrubs. Farther north, in the Chatkal'sk range, following herds of Siberian 
ibex in winter, the predator descends on sunny slopes from heights of 
2,000 to 2,500 m above sea level to below the level of tall junipers 
(Korelov, 1956). 

In Talassk Alatau the ounce inhabits areas between 1,800 and 3,500 m 
above sea level. In summer, following goats [ibex] ascending to escape from 
blood-sucking flies and heat, it also ascends to subalpine or alpine levels; 

226 in winter, chasing goats, it descends along gorges, for example penetrating 
far into Aksu canyon, and visits practically all zones of these mountains. 
In the Aksu-Dzhebagly preserve the snow leopard is not rare in spring and 
early summer at lower levels along river beds and in the tree-shmb belt along 
slopes of gorges, since pregnant females or mothers with young goats, arkhar 
sheep, and roe deer which it hunts are found there at that time (Shul'pin, 
1948; F.D. Shaposhnikov, 1956; V.V. Shevchenko). In the Karatau, the ounce 
is confined to sites of maximum altitudes (1,500 to 2,000 m above sea level), 
namely, the Ker-Dzheilyau where there are many precipitous gorges in which 
wild apple trees, vines, dog rose and other shrubs grow, at places altogether 
impenetrable to man (Antipin, 1953). 

In the Tuyuk gorge of the Kirgiz range adjoining the Talassk Alatau, 
the ounce usually inhabits forestless areas of high mountains, down to the 
upper forest level. There they hunt goats, marmots, and snow cocks 
[Tetraogallus]. Hunting goats at the top of this gorge in May and June 
becomes difficuh and is not without danger. On steep hill slopes the snow 
thaws intensely during the day under solar radiation; these slopes are covered 
either by a very stable and slippery ice crust or a thin and unstable crust. 

227 Avalanches are common at this time of year. In spring pregnant ungulates 
and mothers with young descend to the coniferous forest zone and dense 
shrubs, seeking shelter there. The leopards trail them and are regularly 
encountered in May and June in spruce forests where they mostly catch 
pregnant goats and their young, a fairly easy quarry. In the Karabaltinsk 
gorge in summer snow leopards sometimes descend from the high mountains 
into the juniper tree belt to hunt for roe deer and marmots (Spangenberg 
and Sudilovskaya, 1954). 

In the Kungei Alatau range the ounce is met with from time to time 
in summer in the spruce forest belt (2,100 to 2,600 m above sea level), later 
in the region of creeping juniper and subalpine meadows (2,600 to 2,800 m), 
and very frequently in the alpine level (up to 3,300 m above sea level) 
(B. Petrov, 1950). In the Trans-Ili Alatau and central Tien Shan snow 


226 Fig. 138. Typical liabitat of snow leopard in the Talassk range. A regular trail 

runs through a pass along the crest of the mountain side. Aksu-Dzhebagly preserve, 
western Tien Shan. May, 1955. Photograph by F.D. Shaposhnikov. 

leopards ascend in summer up to 4,000 m or more, sometimes descending 
in winter almost to "shelves" (1,200 m above sea level). In the Altai the 
ounce is also encountered in the alpine zone and on balds, but only at places 
where ibex live (F.D. Shaposhnikov). As in other regions, here too they 
sometimes descend along river gorges. 

While residing in the upper regioas of high mountains, the ounce selects 
228 sections with small open plateaus, gentle slopes and narrow valleys covered 


with alpine vegetation, alternating with rocky gorges and jumbled rocks and 
debris. Siberian ibex and argali sheep live in such places in summer and 
sometimes in winter, and the population of snow leopard there depends on 
their abundance. In the central Tien Shan, it is confined to high mountain 
plateaus, such as elevated watersheds, to which the predator is attracted by 
the presence of argali sheep. Depending on the snow cover and presence 

227 Fig. 139. Kish-Kaindy gorge (2,700 m above sea level). Permanent habitat of snow 

leopards in Aksu-Dzhebagly preserve. Tracks of ounce (male) visible on snow 
in foreground. Talassk Alatau, western Tien Shan. May, 1955. Photograph by F.D. 



of sun-warmed sites, the distribution of ungulates varies and hence that 
of ounce also. In general, however, in winter the ounce is confined to lower 
levels in mountains than in summer, penetrating the forest belt. Never- 
theless, as mentioned above, not only in winter but also in spring and the 
first half of summer this predator often lives in the tree-shrub zone because 
ungulates are abundant there in these seasons of the year. 

Yet it should be noted that the snow leopard is not a high altitude animal 
everywhere. There are regions where it lives year-round in low mountains 
and montane steppes no more than 600 to 1,500 m above sea level. As in 
high mountains, the predator lives here along rocky gorges, cliffs, and rock 
outcrops, since goats and argali sheep prefer these habitats. At a height of 
600 to 1,000 m above sea level it is therefore quite common year-round in 
the spurs of the Dzhungarsk Alatau, i.e., in the Katu and Aktau mountains, 
and also in the Altynemel, Chulak, and Matai (1,000 to 1,500 m above sea 
level). Neither are they rare in the low spurs of the Trans-Ili Alatau and 
Ketmensk — the Malyi and Bol'shoi Bogut, and the Bartogoe, places covered 
with steppe and desert vegetation. All these hills abound in rocky outcrops 
among which goats live regularly, but arkhar sheep are rarer. 

From the foregoing description of the habitats of the ounce, one may 
conclude that this animal is not only an "alpine" animal, as commonly 
assumed, but also a "rock" species to a certain extent. Rock cliffs usually 
contain less snow and hiding in them when threatened is much easier; hence 
goats and rarely argali sheep live in such places. However, the absence of 
snow or a large accumulation on rocks and mountain crests gives this 
predator the advantage of more or less free movement during its hunting. 

Direct observations have revealed that the ounce is poorly adapted to 
moving over deep, loose snow. In areas of loose snow they regularly take 
advantage of beaten trails. When loose snow abounds everywhere the 
229 predator moves along sun-warmed slopes, encouraged to do so because 
ungulates also move there. Thin unstable ice crust can interfere with an 
ounce's hunt. Therefore, in winters of deep snow cover on mountains, ounces 
begin to starve and at these times approach villages and attack domestic 
animals more often than normal. To what extent deep snow cover impedes 
ounce movement can be judged from the fact that two top-notch hunters, 
encountering this predator on an open slope covered with deep snow, quickly 
overtook and captured it by throwing a chapan (top coat) over it and bundling 
it up. 

In rocks it is quite easy for an ounce to conceal itself since it has a 
fairly good protective coloration. Detecting an ounce hiding or creeping 
among rocks is very difficult. Furthermore, the snow leopard is well adapted 
to the severe climatic conditions prevailing in the mountains of Middle and 
Central Asia. Of all the big cats this species boasts the longest and most 


228 Fig. 140. Cirque of Bol 'slioi Alma-Atinsk Lake on tlie northern slope of tlie Tians- 

Ili Alatau, iiabitat of snow leopard. Eastern Tien Shan, 2,650 m above sea level. 
January, 1962. Photograph by A.A. Sludskii. 

dense hair; in addition there is hardly any difference in density of hair 
between winter and summer coats. In the Himalayas ounces have been 
sighted at heights of 5,400 to 6,000 rn (Lydekker, 1924) and are usually 
not encountered below heights of 2,000 to 2,500 m. In summer they are 
seldom seen below heights of 4,(Ю0 to 4,500 m (Zherdon [= Jerdon], 1874*; 
Pocock, 1939). 

Food. The main prey of the ounce almost everywhere year-round is 
the Siberian ibex, and more rarely argali sheep, roe deer, young hog, and 
other small ungulates. In the Pamir it mostly feeds on roe deer (Rozanov, 
1935; Flerov, 1935; Meklenburtsev, 1948; Egorov, 1955), attacking argali 
sheep occasionally. In the eastern Pamir ounces rarely attack wild sheep, 
especially adults. In the Gisser range, in addition to ibex, ounces hunt 
marmots and snow cocks (Leviev, 1939). In the central ranges of the Tien 

■'Not in Literature Cited — Sci. Ed. 


Shan snow leopards feed mainly on ibex and argali sheep; quite often they 
catch snow cocks, marmots, and even relict ground squirrels* and mouse- 
like rodents (Shnitnikov, 1936; Kuznetsov, 1948), in addition to roe deer, 
hare, and rock partridge (Kashkarov, 1932). 

In the Talassk Alatau the main prey of the ounce is the ibex (Shul'pin, 
1948; Shaposhnikov, 1956). The remains of argali sheep, roe deer, piglets, 
and infrequently domestic sheep are found there. Sometimes in winter snow 
cocks and rock partridges become prey (Shul'pin, 1948). The above are 
usually caught at night. In the forest belt of Aksu-Dzhebagly preserve 
(Talassk Alatau) a two-year-old bear killed and partly eaten by an ounce 
was found (Fig. 144). Judging from traces of the fight, the bear was taken 
unawares while digging for roots in a rock stream. It tried to save itself 
by climbing onto a rock. The leopard had eaten the rear portion of the body 
and the viscera but had not touched the remainder, or the skin (F.D. 
Shaposhnikov, 1956). Here this predator had been lying in wait in the early 
morning for roe deer approaching on a solonetz. 

229 Fig. 141. Habitat of the snow leopard around Lake Issyk, Trans-lli Alatau, eastern 

Tien Shan, August, 1960. Photograph by V.G. Heptner. 

*Spermophilus rel ictus — Sci. Ed. 


Fig. 142. Lx)w desert mountains, habitat of the ounce in Chuiak mountains, spurs 
of the Dzhungai-sk Alatau, eastern Tien Shan. September, 1949. Photograph by 

MA. Kuz'mina. 

In Kirgizia and Trans-Ili Alatau the main prey of ounce also is ibex 
but quite often roe deer become victims, especially in winter (lonov, 1929; 
Sludskii, 1939). At some places in the Trans-Ili Alatau roe deer are primary 
winter food of the ounce (B.A. Beloslyudov). The summer lair of one predator 
was littered with a large number of remains of ibex and two piglets. In 
the Dzhungarsk Alatau one often comes across goats killed by the ounce 
(L. Sviridov). In the low spurs of this range (Altynemel', Chuiak) the 
stomachs of two predators caught in November contained only remains of 
ibex (V. V. Afmogenov), while in other spurs (Aktau and Katutau mountains), 
the ounce feeds mainly on the abundant arkhar sheep, and also catches 
goitered gazelle (Antipin, 1941 and 1953). In the Altai and Sayans the snow 
leopard lives only in those ranges where ibex are common. 

In the Himalayas the ounce attacks ibex, gorals, wild sheep, small species 
of deer, Tibetan hares and marmots, and of birds — pheasants. In Kashmir 
they kill domestic goats and sheep, and also horses (Zherdon [= Jerdon], 
1874*; Pocock, 1939). In the Mongolian Peoples Republic the main prey 
is ibex, more rarely argali sheep, and sometimes snow cocks and rodents. 

*Not in LiteraUire Cited — Sci. Ed. 


In the Trans-Ili Gobi the snow leopard may descend into oasis groves to 
catch kulan and goitered gazelle on their way to water holes (a reported 
attack of this predator on aduh kulan is very unsubstantiated). In the 
Mongolian Peoples Republic, the ounce also attacks domestic cattle but such 
reports are few (Bannikov, 1954). In China, in the region of Lake Kukunor 
[= Qinghai Lake], the remains of goats*, argali sheep, eared pheasants, and 
even pigeons were found around two dens of snow leopards with cubs. 
231 It is quite evident that throughout its range the ounce is almost a true 

stenophage since their major food nearly everywhere is Siberian ibex; argali 
sheep, roe deer, and other animals are eaten only in a few places. 

The ounce hunts ibex of all ages, but more often females and kids. Early 
in summer it catches mainly kids. Some summer feces of this predator were 
analyzed and consisted exclusively of the remains of young goats (Ognev, 
1940; Shul'pin, 1948). Among the remains of 14 goat kids found in the feces 
of ounces in the eastern Pamir, only two contained the horny remains of 

Fig. 143. Remains of argali sheep (Ovis amnion karelini) icilled by snow leopard 

at a salt lick. Aksu-Dzhebagly preserve in the Talassk Alatau, western Tien Shan. 

June, 1953. Photograph by F.D. Shaposhnikov. 

*Not clear that this is meant to refer to domestic goat, but the wild goat, or ibex, does 
not occur around Kukunor. — Sci. Ed. 


hooves of kids aged three or four weeks; the remainder were those of 
newborn kids. Evidently ounces catch kids before they are able to follow 
behind their mother. From June to August they attack from time to time 
even adult goats 4 to 10 years of age (four instances). 

Many ibex are killed by the ounce in winters of heavy snowfall. For 
example, in the winter of 1949/1950 the remains of 25 ibex killed by this 
predator, including 17 males, were found scattered over a stretch of 10 km. 
Ounces generally kill old males weakened during rut in winter. Of 30 goats 
killed by ounces, 8 (27%) were females. Of the males killed, 64% were six 
years or older and only 36% four or five years old (Egorov, 1955). In regions 
where the ounce population is high ibex become scarce or disappear 
altogether (Koksu, Dzhungarsk Alatau — 1946 and 1950; Lugovaya region, 
Kirgiz Alatau— 1946). 

The ounce rarely attacks domestic animals, chiefly sheep and goats while 
grazing on alpine meadows (Kashkarov, 1927; lonov, 1929; Sludskii, 1939; 
and others). Sometimes the predator will sneak into a pen containing a large 
number of animals and kill several but a lone individual is more common. 
For example, in the winter of 1955 an ounce attacked a flock of sheep grazing 
on a winter pasture in the Koilyu valley (central Tien Shan) and killed eight 
sheep. In February, 1952 an ounce living in the Ozerno gorge killed seven 
233 sheep. In November, 1953 snow leopards descended very low along the 
Kazachka River gorge near Alma-Ata, and killed four goats (A.S. 
Galunshchikov). In February, 1957 in the Kosh-Agach region (Altai) one 
broke the roof of a sheep pen at night, but was held at bay by dogs and 
shot. Under exceptional circumstances a snow leopard may kill even big 
animals: in 1927 one killed a young horse (lonov, 1929) and in 1938 a pair 
of ounces killed two colts on the northern slope of the Talassk Alatau 
(Kuznetsov, 1948). In winter, with the formation of a deep snow cover which 
makes hunting difficult, ounces sometimes approach human dwellings and 
attack cattle more often than in summer. In November, 1928 in the Saur 
an ounce invaded a sheep pen one night and killed 82 sheep. On Febraary 
11, 1937 a pair of young predators sneaked into a shed and killed a pig 
(Alma-Arasan, 20 km from Alma-Ata; Sludskii, 1939). 

In zoological gardens the daily ration for an ounce is 2.2 kg of meat 
(Obukhov and Shakhnazarov, 1949); in the wild it probably consumes no 
more than 3.0 kg of meat at one time. 

Home range. The dimensions of the individual territory of the ounce 
have not been accurately established. It often travels such long distances 
for a single prey that it is not possible to trace daily movement. While 
bypassing grazing ground of ibex or descending from the higher mountain 
belt into a low-lying gorge, the animal invariably sticks to the same trail, 
which usually runs along a ridge or along a stream or brook. Migrations 


232 Fig. 144. Remains of two-year-old bear {Urstis arctos isabellinus) torn apart by 

two ounces. Aksu-Dzliebagly presei"ve in tlie Talassk Alatau, western Tien Shan. 
May, 1953. Photograph by F.D. Shaposhnikov. 

of the ounce were studied for several consecutive years in a section of one 
gorge near Alma-Ata. From fresh tracks after summer rains or in fresh winter 
snow it was established that the ounce traveled from the eastern slope of 
the mountain into the gorge, moved to another mountain, and returned some 
days later by the same route (lonov, 1929). 

In Aksu-Dzhebagly preserve (Talassk Alatau) adult animals confine 
themselves in winter to a particular hunting territory. The snow leopard 
regularly covers its hunting area, visiting the winter grazing grounds and 
shelters of wild ungulates known to it. The distance covered during these 
visits is considerable and hence a predator is generally seen a second time 
in a given site only after seven or eight days, i.e., fresh tracks appear after 
this lapse of time (F.D. Shaposhnikov, 1956). In November to January, 
1949/1950 in the Tersk Alatau, herds of ibex were confined to definite 
grazing sections and ounces observed there. At one place after a snowfall 
a leopard was regularly seen around an active apiary (Zimina, 1953). Its 
visits were evidently due to scarcity of food. 


232 Fig. 145. Snow leopard chasing ibex (Capra sibirica). Broken line in upper right 

part of photograph denotes ounce track, and in left bottom part tracks of a group 
of mountain goats which had crossed that place 10 minutes before. Dark spots 
on slope are bushes of dwarf juniper {Juniperns). Talassk Alatau in Aksu-Dzhebagly 
presei-ve, western Tien Shan. February, 1961. Photograph by E.N. Mat>'ushkin. 

234 The present population density of snow leopard can be estimated from 

the fact that around the 1950 's about six to eight animals were counted 
in an area of about 40,000 hectares in the Aksu and Dzhebagly River valleys 
of Aksu-Dzhebagly preserve (Talassk Alatau) (F.D. Shaposhnikov, 1956). 
Burrows and shelters. The ounce usually sets up a den in a cave or 
crevice among rock piles under an overhanging cliff or some similar place 
where concealment during the day is guaranteed. Occasionally they rest in 
daylight hours in a sparse spruce forest, among juniper shrubs, in talus slopes, 
or individual rocks. In the Kirgiz Alatau the ounce has reportedly utilized 
the large nests of black vuhures [Aegypiiis monachus] situated among low 
junipers for a diurnal bed. The amount of the predator's fur found in such 
nests indicates that it must spend a significant part of the day in them 
(Spangenberg and Sudilovskaya, 1954). The ounce often uses the same lair 
for several consecutive years. Dens with cubs are always littered with bones 
of ibex and other ungulates. 


233 Fig. 146. Track of snow leopard (male) in packed snow. Aksu-Dzhebagly preserve 

in Talassk Alatau, 3,000 m above sea level. Western Tien Shan. May, 1955. 
Photograph by F.D. Shaposhnikov. 

Daily activity and behavior. The ounce becomes active at twilight, 
hunting before sunset and again at dawn. It rarely hunts during the day. An 
ounce was observed chasing argali sheep at 9.00 a.m., and twice roe deer 
were found that had been killed by these predators at midday (N.A. 
Yanushko). The ounce has often been encountered during the day but was 
usually sleeping or resting on a rock. In the soutnem part of its range, in 
the Himalayas, the predator emerges to hunt even before sunset (Pocock, 
1939). Captive animals lie in a corner of their cage throughout the day and 
begin to move only after the onset of twilight (Fig. 145). 

The ounce hunts by two methods: either the predator creeps toward its 


prey from behind cover or lies in wait near trails leading to a water hole 
or salt lick, hiding on a cliff, in a precipice, or between rocks. When the 
prey is just a few tens of meters away, the snow leopard overtakes it in 
huge bounds of 6.0 or 7.0 m and attacks it quickly. ^^ If it does not catch 
the prey in the first attempt, it usually desists after a few leaps. Such a method 
of hunting is illustrated by the following observation. "In August, descending 
into the valley from Tom' pass, I stopped for the night. The sun had already 
set and it was twilight in the gorge. From all sides the calls of snow cocks 
could be heard. Looking at the opposite side of the gorge, I saw a herd of 
ibex grazing quietly a few hundred meters from where I was standing, and 
two ounces, a large one and a small one, evidently a mother and cub, the 
animals, like shadows, crept between the rocks, using the smallest as cover. 
On coming close to the herd, the adult ounce with an immense leap of 6.0 m 
jumped at an animal but missed. After two more bounds after the escaping 
ibex it returned and went away along the gorge accompanied by the cub" 
(Sludskii, 1939). Reports that the bounds of ounce sometimes measure up 
to 15 m (lonov, 1929) are poorly supported. 

Having caught an ibex or roe deer, an ounce throws its prey to the ground 
and, pinning it with the forepaws, rips open the abdomen or throat with 
its teeth. "An eight -year-old goat which died almost before my eyes had 
deep wounds on both sides of its neck, inflicted by the claws of the predator. 
Death was due, however, to the deep wounds in the abdomen through which 
the ounce pulled out the viscera with its teeth" (Egorov, 1955). In addition 
to concealment, the ounce catches prey from an ambush, waylaying them 
from a cliff or among rocks. In the early spring of 1955 in Aksu gorge of 
the Talassk Alatau, an ounce was seen lying in ambush on a large stone 
overlooking the trail along which female ibex went to a water hole 
(Shaposhnikov, 1956). Judging from tracks in the Dzhungarsk Alatau the 
ounce often attacks ibex from an ambush, leaping from above (A. Sviridov). 
Sometimes this predator attempts to chase its quarry. Thus, in the Dzhebagly 
range ounce tracks behind a female arkhar sheep were found for a distance 
of one kilometer (Shaposhnikov, 1956). In mid-June, 1952, at a height of 
235 about 4,000 m above sea level, an ounce was seen frightening argali sheep 
and chasing them for some time (N.A. Yanushko). Unlike the wolf, during 
a hunt the ounce does not attempt to kill several animals at one time and 
is usually satisfied with one. 

At the end of summer, autumn, and in early winter, the snow leopard 
often hunts in families of two to six animals, more often two or three. For 

^^According to Pocock (1939), as an adaptation to hunting by concealment in rocks, 
the eyes of ounce are set so high that while creeping up to a quarry the predator can see 
it without liftins its head. 


example, in the early winter of 1954 in the Aksu River canyon (Talassk 
Alatau), five ounces belonging to a single family were found attacking a 
young hog. The predators fell on the prey almost simultaneously and quickly 
tore it to bits (F.D. Shaposhnikov, 1956). 

Having killed its prey, the leopard generally shifts it beneath a juniper 
tree or rock before feeding. The remains of a prey are usually left behind, 
and are consumed by vultures, bears, or wolves. Only on occasion does the 
predator stay near its kill and chase vultures away. 

The ounce is not a cautious animal and on noticing or encountering 
man does not hurry to cover. In the early morning of April 24, 1954 in Kish- 
Kaindy gorge (Aksu-Dzhebagly preserve in Talassk Alatau), a preserve 

Fig. 147. 1 rack-s of a female snow leopard on slushy snow. Pass toward Aksu River 

(3,200 m above sea level) in Aksu-Dzhebagly preserve. Talassk Alatau, western 

Tien Shan. May, 1955. Photograph by F.D. Shaposhnikov. 


employee heard his dogs barking. Rising from his seat he saw an adult ounce 
just a few meters away. The animal stood in a tense pose with its tail raised 
and the forepaws pressed firmly to the ground. On noticing the man, it turned 
toward him and snarled loudly. The animal was shot and proved to be a 
female (F. Shaposhnikov, 1956). 

At the same time, the ounce is extremely timid toward humans and 
even when wounded will attack the hunter only in exceptional circumstances. 
Animals which fall into traps are usually taken alive by hunters (see below). 
The timidity of this predator may be judged by the following examples. Late 
in summer an ounce entered a sheep pen high in the mountains on a 
dzhailyau*. On hearing the commotion in the pen, the shepherdess rushed 
in and, catching the animal by its tail, with shouts began to drag it away 
from the wounded sheep. The ounce did not attempt to protect itself and 
was killed by shepherds who had arrived on the scene. In the Kara-Kurdzhur 
region of the Tien Shan dogs once attacked an ounce approaching cattle. 
Taking advantage of the attack, a young shepherd threw his chapati [coat] 
over it and bound the predator (Voinov, 1956). Instances are known of a 
captured adult ounce which allowed itself to be stroked after several days 
and even permitted itself to be approached inside its cages. Taken young, 
ounces become fully tame in a very short while. 

236 Fig. 148. Trail of an ounce in loose snow in Aksu Dzhebagly preserve. Talassk 

Alatau, western Tien Shan. February, 1961. Photograph by G. Kuznet.sov. 

''Kazakh word meaning ''high pasture" — Sci. Ed. 


236 Only two instances are known of ounce attacks on humans. On July 
12, 1940, in Maloalmaatinsk gorge near Alma-Ata, an ounce attacked two 
men during the day and inflicted serious injuries on both. Investigations after 
the animal was shot established that it was rabid. In the second case, not 
far from Alma-Ata, an old, toothless, highly emaciated animal jumped from 
a rock onto a passerby in winter. It was stunned by a blow from a stick, 
tied up, and carried to the village (A.A. Sludskii). 

Seaso?iaI migrations and transgressions. These are poorly known. 
Regular vertical movements while following the trail of ungulates are clearly 
known. In summer part of the population moves into the subalpine and alpine 
belts and in winter, with the onset of deep snow, moves into the forest zone 
and even farther below (see "Habitat"). 

An instance is known of an ounce from the southern part of Lake Telets 
ascending into the upper Kyga and reaching Erinat in the Bol'shoi Abakan 
basin (Kolosov, 1939). Such migrations are evidently associated with a search 
for sites rich in game. In looking for food, the ounce enters the Karatau 
and the spurs of the Trans-Ili and Dzhungarsk Alatau. 

From time to time in search of prey or to escape from deep, loose snow 
cover, the snow leopard descends from the mountains and occurs not only 
in foothills, but also in adjoining plains. In February, 1954, a leopard was 
killed in the foothill steppes (about 700 m above sea level) at Sartoga in 
the Uigursk region of Alma-Ata district (Anurov, 1954). In January, 1958, 
another was caught on the plains near the mountains in the Sairamsk region 
of Chimkent district; the animal was lying among reeds by the side of a 
small lake. In March of the same year three ounces (adult female and two 
cubs) were killed among reeds in the floodplain of the Issychka River (about 
600 m above sea level), roughly 60 km east of Alma-Ata. The animals had 
followed hogs migrating from the mountains because the latter had very 
deep snow cover. In July of the same year an ounce was caught at Dzhaison 
in the Chuisk region of Dzhembul district, 80 km from the nearest mountains. 
It lived in a hilly area and was killed while attacking sheep (Tleukulov, 

In the winter of 1957/1958 an adult ounce was found very far away 
from its regular habitat and caught in a hilly area on the northern shore of 

237 [Lake] Balkhash, 60 km east of the city of Balkhash. The predator could 
have penetrated there from the west through the Chu-Ilisk mountains or from 
the east from the Dzhungarsk Alatau. In either case it traveled more than 
600 km from the nearest mountains (I.G. Shubin). The appearance of ounce 
in very low montane zones, in desert plains, and in oases groves, where 
they are confined to rocks, has been recorded in the uninhabited regions 
of the Trans-Altai Gobi in the Mongolian Peoples Republic (Murzev, 1948). 
In this same country, in the Nemegetu depression of the southern Gobi, the 


Fig. 149. Single track of a snow leopard and its tracks in different gaits (walking 

and bounding). Aksu-Dzhebagly preserve, Talassk Alatau, western Tien Shan. 

February, 1961. Sketch by E.N. Matyushkin. 

ounce lives on low argillaceous and sandy cliffs rising to heights of only 
100 m (Efremov, 1956). 

Reproduction. This is not well studied. According to observations in 
zoological gardens, where this predator sometimes breeds, sexual maturity 
in the female sets in at the age of two to three years. In captive animals 
estrus occurs from March to May. In Uzbekistan estrus in leopards occurs 
from the end of February to March (P.F. Ryplya). During the breeding 
periods with the onset of twilight characteristic loud cries similar to bass 
meowing are emitted by the ounce, generally while standing on projecting 
rocks. The duration of gestation is 93 to 110 days (Dzhennison, 1929*; 
Shereshevskii, 1940). 

In the central Tien Shan young are born around mid-May. A den with 
newborn kittens was found on May 15, 1913. Sometimes litters are seen even 
later. Five litters in June and one in July have been recorded (Shnitnikov, 
1936). In the Narynkol region, in the central Tien Shan, however, a female 
pregnant with two fetuses was caught at the end of March, 1957 (D.I. 
Bibikov). In Kirgizia whelping occurs in May and June (Kuznetsov, 1948; 

*Not in Literature Cited — Sci. Ed. 


Berens, 1957). Early in June, 1955, in the central Tien Shan a litter of five 
blind kittens was found. A den was discovered among rocks only 500 m 
away from a camp of geologists (V.M. Katkov). In June, 1949, a female 
was caught with nurslings in the spurs of the Trans-Ili Alatau. In the Talassk 
Alatau, in Chepta gorge, ounce kittens the size of domestic cats were found 
at the end of JMay (Shul'pin, 1948). In these same mountains, in a forest 
of juniper trees at a height of 1,800 m above sea level, on April 24, 1954 
a pregnant female with one well-developed fetus was caught; the body length 
of the fetus (female) was 22 cm, tail 12 cm, and weight 393 g. The fetus 
already had well-developed pigmented pelage. The mammary glands of the 
captured female were enlarged (F.D. Shaposhnikov, 1956; V.V. Shevchenko). 
In this case parturition would have occurred at the end of April or in the 
first few days of May. A litter of two blind kittens was collected in the 

238 Garm region in Tadzhikistan on May 7, 1940. In the Mongolian Peoples 
Republic ounce kittens — two, three, and rarely one — are born in May and 
early June. By the end of July kittens are one-third the size of adult animals 
(Bannikov, 1954). 

Judging from the dates of encounter given above, kittens are usually 
seen in May in the northern half of the range; more rarely parturition takes 
place in the second half of April or in June. Reports referring to the 
appearance of kittens in the Narynkol region (central Tien Shan) in March 
and eariy April, and in Kirgizia in early April, require confirmation (Sludskii, 
1939; D.P. Dement 'ev et al., 1956). 

A litter may consist of one to five cubs. Even larger litters are possible 
in exceptional circumstances; litters of seven cubs have been recorded 
(October, 1954; Dzhety-Oguzsk region, Issyk-Kul district; Berens, 1957). 
Two or three cubs are more common. In the central Tien Shan the ounce 
asually produces two cubs, generally a male and a female. A litter of three 
kjttens with a single male has been observed only once (Shnitnikov, 1936). 
In Kirgizia an ounce may produce two to five cubs, more otten two or three 
(Kuznetsov, 1948; D.P. Dement'ev et al., 1956; Berens, 1957). In the Talassk 
Alatau this cat gives birth to two or three kittens per litter (Shul'pin, 1948). 
Of seven litters discovered in the Trans-Ili Alatau, one contained a lone cub, 
four had two, one had three, and the seventh had five (A.A. Sludskii). In 
the southern Altai families of ounce consisting of two or three animals are 
observed repeatedly (Selevin, 1929). In the Kukunor district of China two 
dens were found, each housing one ounce and a single cub, and in Tibet 
a litter consisting of three cubs was discovered (Pocock, 1939). Among 
ounces breeding in zoological gardens two to four cubs are common 
(Shereshevskii, 1940). 

239 Growth development and molt. Cubs are bom helpless, blind, and with 
their ear passages closed. Measurements of blind cubs (May 7, vicinity of 


238 Fig. 150. Fetus of a snow leopard a few hours before birth. Aksu-Dzhebagly preserve. 

Talassk Alatau, western Tien Shan, April 24, 1955. Photograph by F.D. Sliaposhnikov. 

Garni) are as follows: male — body length 29.0 cm, tail 15.5 cm, and weight 
430 g; female 28.0, 15.5 cm, and 460 g respectively. One male bom in 
a zoological garden had a body length of 237 mm, tail length of 168 mm, 
length of hind foot 54 mm, height of ears 18 mm, and weight 440 g; the 
corresponding measurements for a female were: 230 mm, 181 mm, 53 mm, 
17 mm, and 495 g (Bree, 1965). The pelage is very dense but quite bristly. 
The main shade of the coat on the back in light brown, especially in the 
region of the shoulders and croup. The coat throughout the upper part of 
the body is mottled with dark brown spots about 1.0 cm in diameter. On 
the neck and withers these spots are even smaller. On the croup the spots 
fuse into two broad longitudinal stripes between which some five or six spots 
are scattered. Around the lip, on the cheeks, and around the whiskers, dark. 


almost black markings occur. The entire underside of the body, commencing 
from the lower lip and up to the genitalia is creamy -white. Around the upper 
part of the neck a^ingle row of dark spots forms a neck band. Rarely, spots 
the size of peas are seen on the upper part of the abdomen. The outer surface 
of the tail has dark brown spots. Both the fore- and hind limbs are light 
in color. 

Even in early July the young begin to trail behind their mother. A female 
and two young ounces were encountered in Mynzhilka region (3,000 m 
above sea level), 20 km from Alma-Ata on July 10, 1937. In July, 1949 
another female with two cubs was met with at Aksai. During July and later 
cubs play together in the mornings just like domestic kittens. Even while 
suckling, cubs take part in the hunt, following their mother. In November, 
1948, at Bartoga (Trans-Ili Alatau) a family consisting of a female with 
five young was observed hunting ibex. The young were half the size of 
the adult. At Lake Iskanderkul' in Uzbekistan at the end of February, 1953, 
a female was caught and the two cubs with her were only slightly smaller 
than she (Ishunin, 1961). The young remain with the mother until she comes 
into estrus again. The male plays no role in the raising of cubs. 

Longevity of the ounce is not known. Some have survived for 15 years 
(Pidoplichko, 1956). 

\4olt has not been studied. A gestating female caught in the Talassk 
Alatau on April 24, 1954, had not yet undergone molt (F.D. Shaposhnikov, 
1956). Ounces molt twice a year. The summer coat differs very little from 
the winter coat in density and length, and this difference is less than in 
other cats. 

Enemies, diseases, parasites, mortality, and competitors. Apart from 
wolf, the ounce evidently has no enemies. Wolf and leopard are serious 

Fig. 151. Snow leopard cub, 1.5 months old. Vil'nyus Zoological Garden. 
Photograph by B.B. Marm. 


competitors in the southern parts of the range. Diseases suffered by the snow 
leopard have not been studied thoroughly. Instances of rabies are known 
among wild ounces (see above). In zoological gardens young animals quite 
often die from infectious enteritis, a highly contagious disease with a very 
severe course; mortality may be as high as 70 to 80% (Tsvetaeva,1949). 
Mites, HaemaphysaUs warbiirtoni, were found in an ounce caught in the 
vicinity of Alma-Ata on October 29, 1955 (G.B. Ushakova). 

Population dynamics. In some years the ounce population is observed 
to increase sharply in one or another region. Comparatively large numbers 
were sighted in 1946 in the region of Lugovaya station, Kirgiz Alatau, 

240 and in the Gvardeisk region, Dzhungarsk Alatau. In the latter region ounces 
were numerous in the winter of 1949/1950. In the winter of 1952/1953 an 
increase in snow leopard population occurred in Kaskelensk region of the 
Trans-Ili Alatau mountains and in the Taldy-Kurgansk region of the 
Dzhungarsk Alatau; in the latter region a simultaneous rise in the lynx 
population was also noted. In the following winter of 1953/1954 the ounce 
and lynx population increased considerably again in the Dzhungarsk Alatau 
and even in the Panfilovsk region (V.S. Korovets; M.S. Kuznetsov; and 
others). It is possible that this increment in population of both predators 
was due to an abundance of ibex in these areas. In general, factors controlling 
changes in the population of the ounce are not known. In the 1940's to 
the 1950's a reduction in hunting led to an increase in snow leopard stocks 

Field cliaracteristics. The ounce is a large cat of smoky-brownish-gray 
color mottled with large, annular, dark brown spots. The tail length exceeds 
three-fourths of the body length; it is densely furred and appears even longer 
and thicker. Tracks are large, rounded, and without claw impressions. 
Remains of ibex killed by the ounce are usually seen close to steep cliffs 
and rock piles, under juniper, and in other such sites; wolves kill them only 
on gentle slopes, far away from rock outcrops. 

The remains of prey of the ounce and wolf differ considerably in the 
following respects. In the skull of an ibex or arkhar sheep killed by ounce, 
the nasal bones and orbits remain undamaged. Around the skeleton of an 

241 ibex killed by the ounce invariably one finds large bits of skin. The wolf, 
contrarily, generally punctures the nasal region of the skull, cmshes the 
orbits, punctures the lower part, and frequently enlarges the foramen magnum 
in four- to five-year-old arkhar sheep and ibex to extract the brain. The 
wolf strips the skin into small pieces or eat it completely. Even if a wolf 
has scavenged the remains of the leopard's prey, the fact of the latter 's 
kill can still be ascertained from the large scraps of skin scattered around. 
When the death of an ibex or arkhar sheep is not due to these two predators, 
vultures quickly find the carcass and having eaten the carrion, turn the skin 


240 Fig. 152. Snow leopard on cliff in the Trans-Ili Alatau, eastern Tien Shan. 

Photograph by M. Galkin. 

inside out; they do not tear it to pieces as done by the ounce (Egorov, 1955). 

Practical Significance 

The ounce is of little importance to the fur industry. In the 1950's to the 
1960's only a few tens of skins were tanned everywhere. Consequently their 
tanned value was very low, on the average about 3 rubles. The main regions 
of ounce hunting are Tadzhikistan and Kirgizia. However not long ago in 
Kirgizia, up to 60 skins were prepared annually. Thus, in 1925/1926, 37 


skins were prepared, 1926/1927—56 skins; 1927/1928—20 skins; 
1928/1929—37, 1931—53, 1936—54, 1937—45, 1938—31, 1939—45, 
1940—27, 1942—16, 1943—24, and 1944—13. In the 1950's and 1960's, 
6 to 18 skins were tanned annually in Kirgizia. 

In Kazakhstan in recent years fewer than 10 ounce skins have been 
tanned. A larger number were tanned there formerly. In 1926/1927, 8 skins 
were prepared, 1928/1929—66, 1933—4, 1934—15, 1935—20, 1936—3, 
1937 — 3, 1938 — 13, and 1939 — 14. More skins of snow leopard were 
prepared in Tadzhikistan in the 1950's: 1951 — 16, 1952—8, 1953—17, 
1954—27, 1956—35, 1957—33, and 1958—53. 

In Uzbekistan over the last 10 years, 3 to 8 skins were tanned annually, 
mainly from Surkhan-Dar'insk regions: 1951 — 5, 1952 — 6, 1953 — 6, 
1955 — 6, 1956 — 3, and 1957 — 3. It is thought that presently about two- 
thirds of the skins of snow leopard remain with the people. Some lone ounces 
have been caught in Altai and the Sayans. 

For the Soviet Union as a whole, 118 "leopard [barsy skins were 
tanned in 1928/1929; of these, 8 were prepared by the Siberian group of 
firms, and 110 by the Middle Asian group (Kogan, 1931). This number 
also included skins of common "leopard" [leopard] and cheetah, comprising 
about 20 skins. In the Soviet Union in the 1920's some 100 skins of ounce 
were tanned annually: 1926/1927—70 skins, 1927/1928—30, 
1928/1929—120, 1931—82, 1953—29, 1954—52, and 1956—120 
(Kuznetsov, 1932). 

In the last century ounce hunting was intensified and in the Semirech'e 
district up to 140 caught annually. The magnitude of hunting can be judged 
from the following data on dressed skins (Silant'ev, 1898; Shostak, 1927): 
1884—83, 1885—29; 1886—142, 1890—37, 1891—54, 1892—106, 
1893—47, 1894—74, 1895—36, 1897—58, 1898—76, 1899—29, 
1900—30, 1901—36, 1903—47, and 1905—70. Of these, the bulk came 
from the Przheval'sk and Pishipeksk sections of Kirgizia. Kazakhstan 
provided 10 to 20 skins (Dzharkent section). 

The annual output of ounce skins from the Soviet Union, compared 
with the world output, was very much greater in the past. 

From 1907 to 1910 the annual world output of snow leopard skins was 
placed at 750 to 800. In some years up to 500 leopard skins passed through 
Nizhnegorod market, while the world output was estimated at 1,000 (Zhitkov, 
1928; Kuznetsov, 1932). Thus Russia formerly contributed up to 50% of 
the world output of ounce skins. Mongolia produced 80 skins in 1908, 40 
in 1927, 40 again in 1934, and 15 to 25 annually until 1944 (Bannikov, 1954). 
242 All the 120 leopard skins put up for sale in 1956 at the Leningrad auction 
were sold to the USA, at an average price of $11.25 per skin.These skins 
were mainly used in making carpets and fur coats for women. 


Early in the present century methods of catching live ounces were 
developed in Kirgizia, and catches either exported or placed in Soviet 
zoological gardens. Catching ounces alive is practiced at present in 
Kazakhstan, Kirgizia, and Tadzhikistan. Live animals are handled by 
Zootsenter. From one, the Issyk-Kul' district (Kirgizia) alone, this 
organization in the 1950's received 50 ounces in three years (Berens, 1957), 
while 14 live animals were captured in the first half of 1958 in all of Kirgizia. 
During the 1940's and 1950's the Przheval'sk branch of Zootsentr in Kirgizia 
received 375 live leopards, or an average of 18 animals per year. The 
Tadzhikistan branch of Zootsentr, located in the eastern Pamir, received 
50 leopards in 10 years (1950's). In individual years the organization handled 
up to 10 animals. 

In the world market the ounce is always in great demand and expensive. 
Within the Soviet Union a live animal is priced at several hundred rubles. 
Though the snow leopard became known in Europe even at the end of the 
eighteenth century, they first became acquainted with a live animal only 
in 1872 when Governor- General Kaufmann sent a pair of young animals 
from Turkestan to the Moscow Zoological Garden. 

The damage caused by these predators to animal husbandry in 
Kazakhstan is of no practical significance. In regions where livestock 

242 Fig. 153. Snow leopard taking refuge in a tree. Vaiciiana gorge, Pamir. Photograph 

by G. Lipskerov. 


are many and the ounce common, no more than one or two attacks have 
been reported. Mainly stray sheep and goats become victims (Sludskii, 1939). 
They cause little damage to stock even in Kirgizia where the predator is 
particularly common (Kuznetsov, 1948; D.P. Dement'ev et al., 1956). They 
rarely attack man, even when wounded. Only two attacks are known — 
one by a rabid ounce and another by an extremely emaciated old animal 
(see above). 

Live ounces are caught by placing large traps woven from sheep's wool 
or loops of steel cable along their trails. Loops are usually placed together 
with a small trap with weak springs. Having set up the trap, the loop is placed 
on top of it and the free end tied to a spring on a strong and resilient pole. 
An ounce having stepped into a trap, simultaneously is grabbed by the loop. 
The latter is quickly retracted by the spring, hoisting the animal into the 
air by its paws in such a way that its hind feet can still touch the ground. 
243 To ensure that the loop does not injure the animal's foot, a restrictor is 
first attached which prevents excessive tightening. An ounce on falling into 
a trap usually executes one or two leaps, then sits quietly. It cannot escape 
even if only one or two digits are caught in the trap. 

Fig. 154. With a captured snow leopard. Tien Slian. March, 1959. Photograph 

by A. Grecher. 


The ounce is removed alive from the trap or loop in the following 
manner: one hunter stands in front of the animal and distracts it while another 
approaches from the rear and throws an overcoat or chapati over it in order 
to catch it by the head. The paws are then bound together and tied down, 
a stick is placed between the teeth (tied together), after which the animal 
is released from the trap or loop. Sometimes a trapped animal is noosed 
at the neck, one paw slipped into a loop tied to a beam ("ikryuk"), and 
then the animal tnissed. Often hunters approaching a captured animal adopt 
a different procedure. One man moves forward to distract the ounce or places 
a stick in its mouth, while another at an appropriate moment catches its 
tail and drags the body toward himself with force. The head of the dragged 
animal is quickly covered with an overcoat, padded jacket, or a thin felt 
material made of sheep's wool so as to temporarily blind it. A blinded animal 
quiets down and can be tied up easily. Bound and gagged, the animal is 
then strapped to the saddle of a horse, carried on the back (young animal), 
or pulled on a sledge of branches to the camp and placed in a temporary cage. 

At present, several hunters are known who have successfully trapped 
ounces by the methods described above, among them a widely known hunter 
living in the Issyk-Kul' district (Kirgizia) v.'ho has trapped up to 30 of these 
cats (Berens, 1957), and another famous leopard catcher who caught 60 
in Tadzhikistan (in Murgab region). From time to time ounces fall into traps 
244 intended for ibex or argali sheep. On these occasions they are shot. After 
the first snowfall ounces are sometimes hunted with the help of dogs; 
moreover, at the beginning of this century they were sometimes poisoned 
with strychnine. 

In the standard fur terminology used throughout the Soviet Union ounce 
pelts are called "leopard" [bars]. They go into carpets, sledge covers, 
women's overcoats, and more rarely fur coats, collars, etc. 

For a long time, due to misunderstanding, the ounce was considered 
a dangerous and harmful predator and hunting thus permitted year-round 
by any method. Rewards were even offered for dead animals. In reality, 
as stated before, the harm done to livestock and game is insignificant, and 
the cat poses no danger whatsoever to man. The ounce is, in fact, a valuable 
treasure in our mountains and of great scientific interest as a highly 
specialized wild cat. In the world market live ounces are always in demand, 
and their sale a profitable item for zoological exports. 

Evidently the indiscriminate destruction of this animal should be banned 
everywhere. Catching ounces should be prohibited year-round in the 
Tarbagatai, Saur, Altai, and Sayans. In Tadzhikistan, Uzbekistan, Kirgizia, 
and Kazakhstan shooting ounces should be banned and catching them for 
the purpose of export only permitted from October through February. 

In 1950 in Kirgizia, the ounce was struck from the list of predators to 


be exterminated. Hunting was banned altogether and only the catch of live 
animals permitted. In this same republic capturing cubs and destroying dens 
with cubs are prohibited. (A.S.) 

244 Genus of Small Cats 

Genus Felis Linnaeus, 1758 

1758.^ Felis Linnaeus. Syst. Nat., 10th ed., vol. 1, p. 41. Domestic cat, 

i.e., Fells libyca Forst. 
1792. Lynx. Kerr. A?iim. Kingd. Cat. Mamm., NN 288-289. Fells lytix 

1821 . Lyiicus. Gray. London Med. Rept., vol. 15, p. 302. Fells lynx Linnaeus. 
1829. Pardina. Каир. Entw. Gesch. Nat. Syst. Europ. Thlerwelt, vol. 1, 

pp. 53 and 57. Fells pardina Temm. = Fells lynx Linnaeus. 
1834. Lynchus. Jardine. Nat. Llbr. Mamm., vol. 4, p. 274. Felix lynx 

1841. Otocolobiis. Brandt. Bull. Sc.Acad., St. Petersb., 1, 9, 38. Fells maniil 

1843. Caracal. Gray. List. Mamm. Brit. Mus., p. 46. Caracal melanotis 

Gray = Fells caracal Schreber. 
1855. Catiis. Fitzinger. Wiss. Popul. Nat. d. Sdiigeth., vol. 1, p. 265. Fells 

catiis Linnaeus (domestic cat). 
1858. Catolyx. Severtsov. Revue. Mag. ZooL, vol. 10, p. 387. Fells chaus 

1858. Prlonallurus. Severtsov. Ibid., p. 387. Fells pardochrous Hodgson 

= Fells bengalensis Kerr. 
1858. Urolynclms. Severtsov. Ibid., p. 389. Felis caracal Schreber. 
1858. Oncoldes. Severtsov. Ibid., p. 390. Given to one of the South 

American cats but applied to euptllura Elliot (Satunin) with a 

reference to Severtsov. 
1867. Cervaria. Gray. Proc. Zool. Sac. London, p. 276. Lyncus pardlnus 

= Fells lynx Linnaeus. 
1898. Oncoldes. Trouessart. Cat. Mamm., vol. 1, p. 357. Nomen 

praeoccupatum (Severtsov, 1858). 
1903. Eucervaria. Palmer. Science, N. S., vol. 17, p. 873. Replacement 

for Cervaria Gray. 

245 1905. Trichaelurus. Satunin. Ezheg. Zool. Muzeya /. Ak. Nauk, vol. 9, p. 

495, St. Petersb. Replacement for Otocolobus Brandt, 
1926. Eremaelurus. Ognev. Ezheg. Zool. Muzeya L Ak. Nauk, vol. 27, 

^Some of the names given were coined for subgenera, but also used as generic names. 


p. 356, St. Petersb. Ei'emaelurus thinobius Ognev = Felis margarita 
Loche. (V.H.) 

Small- and moderate-sized cats; a few species are large. 

The general outward appearance is typical of small cats, i.e., sacral 
region high and stands above shoulder, and back arched forward; more rarely, 
back line straight (see Fig. 30). Legs mostly short or moderate in length, 
as a result of which the animal appears long; a few are long-legged and 
these animals appear nearly squarish. Tail varies in length; in a few species 
very short, not more than one-seventh of body length (trunk with head), 
and does not reach calcaneal joint in standing animal; in most species of 
moderate length, varying from one-third to one-half of body length; and 
in a few species equal or almost equal to body length. 

Head relatively large, short and broad in most, almost spherical in some, 
slightly elongated in a few, and perceptibly so in some, resembling somewhat 
the head of big cats (genas Panthera; puma and clouded leopard). Ears vary 
in size and shape but almost invariably relatively larger than those of big 
cats (Panthera) and ounce (Uncia), and prominently project beyond even 
the udnter coat. Usually they are quite large and pointed but sometimes blunt; 
sometimes they are very large and steeply set; or they may be pushed sideways 
("overhung") and the parietal region between them very broad. In some 
species the hair on the top of the ear is only slightly elongated; sometimes, 
however, the long pointed ear has a well-developed tuft of long hair at the 
tip. Eyes relatively large and very large in most species; pupil usually vertical 
but round in some. Hair coat uniform throughout the body; hair on cheeks 
in a few species long and forms fairly well-developed whiskers. 

The feet in most species are not strong and the paws relatively small; 
in a few big and moderate-sized species, they are strong with powerful, 
broad paws similar to those of ounce and big cats (puma, lynx, and clouded 
leopard). The claws in all species except one (F. planiceps, Kalimantan) 
are wholly retractile and their ends do not project out of the fur. Skin folds 
and interdigital membrane quite variable, indicating various degrees of 
development of this adaptadon. In some species (clouded leopard, marbled 
cat — F. marmorata, and others), interdigital membrane and skin folds over 
claws fully developed, i.e., folds are doubled up, occur on all the digits, 
and the membrane reaches ends of digital cushions (as in most cats and 
ounce; see Fig. 278). In many species the skin fold is the same size on 
each digit in both fore- and hind paws, and covers retracted claws of digits 
II and III from the inside and of digits IV and V from the outside, while 
the interdigital membrane is relatively pooriy developed. In some species 
skin folds cover the claws but the interdigital membrane is almost 
undeveloped (caracal, similar to cheetah; see Fig. 278).^ 

^Description of paw structure here and elsewhere taken from Pocock (1941). 


Pelage usually dense, soft, silky, and relatively long, especially in winter. 
Color highly variable, from wholly monochromatic, often yellow shades, to 
monochromatic with traces of spots, usually on the extremities, of various 
color shades. Spots highly variable in size, ranging from blotchy to annular 
to small, or small fused into large ones. Often these spots form bands or 
fuse, or almost so, into longitudinal and more rarely transverse small stripes. 
A true ''tiger" color does not occur. In some species (clouded leopard, group 
Prionailurus) a white spot occurs on the back of the ear as in some big cats. 
A few species are polyphasic {F. temmincki and F. yagiiarundi); melanistic 
and erythristic individuals occur in many species and are sometimes 
geographically localized (leopard cat and manul). 
246 Teats, two to four pairs. 

Skull of most species relatively large, light in build, and with relatively 
thin bones. Zygomatic arches weak, orbits large and closed or almost closed 
in some species, and cranium large and bulging. Upper line of profile highly 
convex, fairiy steep, and arcuate; interorbital region raised and profile, both 
behind and in front of interorbital region, dips steeply. Skull broad posteriorly; 
zygomatic width may exceed one-half of condylobasal width. General skull 
features viewed dorsally appear elliptical to varying degrees with a slightly 
elongated cranial part. Exceptionally, skull appears rounded. 

Crests and tuberosities absent or only weakly developed in most species; 
sagittal crest totally absent or seen only posteriorly in the form of an 
insignificant elevation; lambdoidal crests better developed but still weak. 
Postorbital constriction weakly defined. Facial part of skull relatively small 
and short, usually shorter than cranial part (reckoned from middle of 
supraorbital processes). Small pit (depression) present between nasal and 
frontal regions. 

In most species the gap between the posterior wall of the glenoid fossa 
(postglenoid process) and the tympanic bulla is very small. In a few the 
bullae are shifted significantly behind, or contrarily, set so far forward that 
the anterior margin of the tympanic bulla falls at the level of the glenoid 
fossa. Tympanic bullae per se very large, relatively much larger than in 
big cats and ounce, and bulging. Usually the entotympanic chamber is larger 
than the ectotympanic; in some species, however, the latter is equal to or 
only slightly smaller than the entotympanic chamber. 

The third upper element (epihyale) of the hyoid apparatus (see Fig. 31) 
is completely ossified, and depressions in front of the jugular foramen 
are absent on the basioccipital part of the occipital bone. Only in a few 
species is the skull relatively long and coarser, with strong zygomatic 
arches, well-developed crests, and a more powerful postorbital constriction, 
giving it the general appearance of the skull of big cats (genus Panthera; 
clouded leopard, F. nebulosa, the skull of which resembles that of puma. 


F. coiicolor; and to a lesser extent several other species, for example 
F. temmincki). 

Dentition in most species is complete although the second upper 
premolar (first in the series) is very small. Some individuals in certain species 
lack this tooth; in other species, often distant from one another, the general 
absence of this tooth is designated a specific characteristic in spite of the 
fact that it is present in some individual animals. The dental formula in 
this genus may there be given as: 

.31 31,^ 
1 , с , pm , m =30 

3 1 2 1 

3 1 '^ I 

or i — , с — , pm — , m — = 28 

3 1 2 1 

As an abnormality a nidimentary second lower molar is sometimes 
seen (lynx; Zoological Museum, Moscow University). Relatively weak 
teeth have also been noted in which the talon of the upper camassial tooth 
was greatly reduced in size and the canines relatively thinner at the base, 
more slender, and relatively longer than in big cats and ounce. Particularly 
long canines (longest in the family) have been recorded for clouded 

Sexual dimoфhism is absent but females average smaller. Age-related 
dimorphism is manifested in several species, mainly monochromatic or 
weakly spotted ones. Юttens have either well-developed (puma) or only 
faint spots throughout the body or only in some parts (extremities). Seasonal 
color dimoфhism is absent or the winter coat perceptibly lighter than the 
summer one (lynx). In most species the winter coat is denser and more 
luxuriant (longer) than the summer coat. 

These are predators that prey mainly small vertebrates, mostly rodents 
and birds. In other words, the quarry is usually much smaller in size than 
M7 the predator. Some large-sized species (puma and lynx) catch hares and 
animals even bigger than themselves (deer). They hunt by ambush. Members 
of the genus inhabit all types of terrain except tundra and occupy highly 
diverse biotopes ranging from high latitudes in mountains to sandy deserts. 

Due to the specialized structure of the hyoid apparatus (see Fig. 31) and 
the poor mobility of the larynx, these animals cannot produce a low "roar- 
growl". Instead their call is very high pitched shriek, resembling that of 
domestic cats, and varies widely in different species ("meowing cats"). 
Well-being and contentment are expressed by purring during both inhalation 
and exhalation. By and large cats produce diverse types of sounds, and hence 
members of the genus Felis also growl, hiss, snort, and produce a peculiar 
snarl. They mainly eat while sitting hunched on the hind legs. The front 


legs and forearms do not touch the ground. Pieces of the prey are usually 
bitten off and they do not tear it with upward jerks of the head (see Fig. 32). 

The geographic distribution of the genus corresponds to that of the 
family but small cats are evidently absent in some regions within the range 
of the family. In some countries (Europe and North America) the range 
has undergone great reduction as a result of extinction during the last century. 

As pointed out earlier (see characteristics of the family), the genus Felis 
s.l. represents a progressive group of cats, the members of which reflect 
most distinctly the main line of specialization of the family. It is the highest 
type of cat, and it may be, of predators in general. It is closely related to 
big cats, Panthera. The snow leopard, genus Uncia, is a direct link between 
Felis and Panthera and a transitional form in the classification adopted 
in this work. 

Nevertheless genus Felis is very diverse and includes forms specialized 
in different directions. Species large in size and adapted to catching relatively 
big animals belong to this group. While preserving the basic diagnostic 
features of the genus, these cats include extreme forms which also exhibit 
a similarity of features with big cats (Panthera) and the ounce. Members 
of genus Felis include forms transitional between highly specialized species 
of small felines (sand cat and manul) and ounce and thus big cats (more 
precisely, the reverse). Examples are the puma and clouded leopard. 

As mentioned several times before, there is no unanimity of opinion 
with regard to the composition of the group of cats considered here under 
the single genus Felis. Some authors divide this group into 12 (Haltenorth, 
1957), 14 (Pocock, 1917), 16 (Weigel, 1961) and even 17 (Mazak, 1965) 
genera. Diversity, undoubtedly considerable, cannot be ignored by adopting 
a single large genus. This simply emphasizes the gradual evolution of 
characteristics in the group and the mutual link between what at first glance 
are dissimilar species or even polar opposites (clouded leopard — sand cat 
and manul) through a series of transitional forms. At the same time such 
a division naturally leads to the recognition of subgenera — a category 
particularly suited for these situations. 

Various Soviet species have been grouped into three subgenera in this 
work. The subgenus Prionailurus usually includes three South Asian species 
which judging from the fourth species inhabiting the Soviet Union (F. 
euptilura), cannot be supported.^ In the subgenus Felis itself different 
zoologists include five or six Euro-Asian and African species, including 
the sand cat from Turkestan, the Arabian Peninsula and North Africa which 
is treated as two separate species (margarita and thinobia). This view cannot 

■'The Amur forest cat produces fertile hybrids with the domestic cat in the Ussuri 
region (see below). 


be accepted. First of all, these are simply two forms of the same species 
(Heptner and Dement 'ev, 1937), and secondly they differ from true Felis 
s. str. in several characters, of which the most important peculiarity is the 
structure of the tympanic bulla. The South African F. nigripes (subgenus 
Microfelis) should evidently be excluded from this subgenus. 
248 The subgenus evidently comprises seven species but its composition 
differs from that generally recognized: three species invariably regarded 
as Felis s. str. — silvestris (including group libyca), chaus and bieti and 
three species usually placed with Prionailurus — viverrinus, bengalensis, 
and rubiginosus. 

F. euptilura usually placed in subspecies bengalensis, also belongs here 
but is treated as an independent species in the present work. Of these seven 
species, three inhabit the Soviet Union. Subgenus Lynx includes three species 
of which the Soviet fauna has two and subgenus Otocolobus, two species, 
both represented in the Soviet fauna. Outside the Soviet Union small groups 
(two or three species) belonging to fairly distinct or identifiable subgenera 
exist. More importantly, some individual species deserve elevation to the 
rank of subgenus (ocelot, serval, flat-headed cat, F.planiceps, jaguarundi, 
clouded leopard puma, and others). 

Applying the same principles as used for evaluating relations between 
genera in the family relations between the subgenera and individual species 
of the Soviet (and in part American) fauna listed above are as follows. Of 
the three subgenera, two (Felis and Otocolobus) may be considered forms 

Subgen ^ 

F. rufao 

F. lynx- 


о group lynx 

oF. manul \ 

I Subgen 
\ Otocolobus 

6F. margarita* 

о group libyca 

° silvestris 

* silvestris 

^ Felis 

F. caracal о oF. chaus 

oF. euptilura 

Fig. 155. Systematic relations between some cats of the genus Felis in the northern 
Holarctic. V.G. Heptner. 


representing one line ("direct") of development (specialization) toward the 
attainment of a completely feline type. Subgenus Fells, however, is less 
specialized than Otocolobus. The third subgenus, Lynx, represents a lateral 
line, highly similar and parallel to the general line of the first two, but not 
attaining the level of specialization oi Otocolobus insofar as skull structure 
is concerned. 

The main feature of subgenus Lytix is the general body build, which 
is very typical and exhibits no variation in the various species of this 
subgenus. These cats stand on long legs and sport short tails; the contour 
of the body and legs therefore approaches the form of a square. The 
subgenera Fells and Otocolobus are characterized by long bodies, long legs, 
and a relatively long tail. The progressive specialization of the skull in these 
two subgenera has reached (in several species) a level far higher than in 
subgenus Ьу?гх, and the highest level in the family (subgenus Otocolobus 
— O. manul). 

Insofar as relations between species are concerned in the main line 
(Fells'^' — Otocolobus) these are defined by a progressive reduction in the 
facial region of the skull (and a corresponding increase in the cranial region), 
a progressive turning of the orbits forward, and a partial alteration in the 
structure of the auditory apparatus. Highly specific adaptations have 
developed against this "phyletic" background (for details, see "Description" 
of the species and particulariy the section ''Systematic Position". 
249 On the whole, our fauna is represented by the following series: F. (F.) 

euptilura (Amur cat) — F. (F.) chaus (jungle cat) — F. (F.) silvestris, group 
silvestrls (European forest cat) — F. (F.) silvestris, group libyca (steppe 
cat) — F. (O.) marganta (sand cat) — F. (O.) manul (manul) (Fig. 155). 

Species of subgenus Lynx (lynx), which exhibit the main feiitures of 
the genus maximally, i.e., structure of body and legs and general degree 
of specialization of the skull (in the direction Fells — Otocolobus), form 
the series: F. (L.) caracal (caracal) — F. (L.) lynx, group lynx (Eurasian lynx) 
— - F. (L.) lynx, group canadensis (American lynx) — F. (L.) rufa (bay lynx 
[bobcat] of America). Specialization (progressive advancement) of the skull 
in this series of forms (species and subspecies) and differences in the skull 
features between the initial and terminal species in the group {caracal — 
rufa) are less than in members of the corresponding main series {Fells — 

The relative level of specialization among species of the subgenus Lynx 
(lynx), compared to that of the main line, is such that the skull of F. (L.) 
caracal (caracal) corresponds, or almost so, to the skull of F. {F.) chaus 
(jungle cat), while the skulls of the two groups of true lynxes, F. {L.) lynx 

* Felix in Russian original — Sci. Ed. 


{lyfvc and canadensis) and F. (L.) rufa (bay lynx), reach only the level of 
F. (F.) silvestris, group libyca, or slightly higher, but not the level of F. (O.) 
margarita (sand cat) (Fig. 155). 

Paleontological data pertaining to the genus are extremely meager. The 
genus Felis, like the genus Panthera, obviously evolved in the Pliocene 
and its differentiation is of recent origin. Moreover, a true lynx, F. QL.) 
brevirostris has been found in the upper Pliocene. This group originated 
earlier than the Early Pleistocene (Villafranchian) — F. (L.) issidoriensis, 
through F. (L.) tellhardi to present-day lynx, appearing in the Upper 
Pleistocene. A distinct species of puma, F. (P.) inexpectatus, is known in 
the Pleistocene of America (Thenius and Hofer, 1961). 

The genus Fells is very rich in species of carnivores, in fact one of the 
richest genera among mammals in general. It contains 29 extant species'^: 
F. bengalensls Kerr, 1792 — A; F. rublgliwsus Geoffrey, 1834 — A; F. 
vlverrlnus Bennett, 1833 — A; F. euptllura Elliot, 1871 — A; F. planlceps 
Vigors and Hors field, 1828 — A; F. marmorata Martin, 1836 — A; F. badla 
Gray, 1874 — A; F. temmlnckll Vigors and Horsfield — A; F. aurata 
Temminck, 1827 — A; F. pardalls Linne, 1766 — SA; F. wledi Schinz, 
1821— SA; F. pardalls Linne, 1766— SA; F. wledl Schinz, 1821— SA; F. 
colocolo Molina, 1782 — SA; F. geoffi-oyl d'Orbigny and Gervais, 1843 — SA; 
F. pardlnoldes Gray, 1867 (tigrlna) — SA; F. gulgna Molina, 1810 — SA; 
F.pajeros Desmarest, 1816 — SA\ F . yaguarundl Lacepede, 1809 — SA CA 
and NA; F. chaus Giildenstaedt, 1776 — A and Af; F. silvestris Schreber, 
1777 — A and E (including F. libyca Forster, 1780 — Af, A, and E); F. bletl 
Milne-Edwards, 1892 — A; F. serval Schreber, 1776 — Af; F. lynx Linnaeus, 
1758 — A, E, and NA; F. rujus Schreber, 1777 — NA; F. caracal Schreber, 
1776 — A and Af; F. nlgrlpes Burchell, 1882 — Af; F. margarita Loche, 
1858 — A and Af; F. manul Pallas, 1776 — A; F. concolor Linne, 1771 — SA, 
CA and Na; F. nebulosa Griffith, 1821— A.-*^ 

Species of this genus comprise 82% of the total number of species of 
the family. Most (17) are from Asia; moreover all, except one (lynx), are 
distributed in the southern half of the mainland. Of these, four are common 
to Africa, two to Europe, one to Europe and North America, and twelve 

''This list is only a catalog and does not represent a specialization sequence. E = Europe, 
A = Asia, Af = Africa, NA = North America, SA = South America, CA = Central America. 

^This list does not include Mayailurus iriomotensis Imaizumi, described in 1967 as 
a new genus and species from Iriomote Island, one of the southernmost islands of the Ryukyu 
chain lying on the latitude of Taiwan. This form is described by the author as coming close 
to the South American F. giiigna ("Oncifelis giiigna"). The absence of specimens and the 
nonavailability of a full description of the species do not permit a positive judgment about 
the position of this cat from Iriomote. It is quite clear, however, that firstly, another new 
genus has no justification from the viewpoint adopted here and, secondly, this cat probably 
belongs to the group Prionailums (it gready resembles F. viverrina and partly F. euptilura). 


solely Asiatic. In South America there are nine species, of which two are 
common to Central and North America and seven exclusively confined to 
the mainland. In number of species Africa stands third with six, of which 
250 four are common to Asia or Asia and Europe, and only two are endemic. 
In North America there are four species, of which one is common to Eurasia, 
two to South and Central America, and only one endemic. In Europe there 
are two species, of which one is common to Asia and North America and 
one to Asia and Africa. 

There are no endemic species in Europe. 

Evidently there are two clearly distinct centers of cladogenesis of 
cats — South America and Asia (southern half). If one takes into account 
the respective number of species of big cats (Panther a) and ounce, the link 
between most species of the family from southern Asia becomes even more 
striking. The center of development of present-day species of the family 
would seem to lie in this part of the globe, probably in the extreme south 
and southeast sectors. 

These are fur-bearing animals of little economic value. Some species 
are large (lynx, but more so puma) and objects of sport. They control rodent 
and hare populations and sometimes even large game. Occasionally they 
harm game populations, primarily through decimation of game birds, and 
sometimes domestic horned animals. 

Six species are represented in the fauna of the USSR: Amur cat, F. 
(Felis) euptilura Elliot, 1871; European wild and steppe cat, F. (Felis) 
silvestj-is Schreber, 1777; jungle cat, F. (Felis) chaus Giild., 1776; caracal, 
F. {Lynx) caracal Schreber, 1776; lynx, F. (Lyfix) lynx Linn., 1758; sand 
cat, F. (Otocolobus) inargarita Loche, 1858; and manul, F. {Otocolobus) 
manul Pall., 1776. 

These species constitute about 21% of the species of the genus and 
about 8.0% of the total number of mammals in the Soviet fauna. 

The range of the genus Felis in the Soviet Union covers all of the forest 
zone, including montane forests in the southern part of the country, the 
semidesert zone (in part steppes), and desert regions. The range has shrunk 
greatly at places in the west in recent years. 

In certain areas (Middle Asia) they are fur-bearing animals of no mean 
consequence. In other regions in the south they hurt commercial hunting 
by decimating game birds. Some species are of vital importance in the live 
animal trade. (V.H.) 



Felis (Felis) euptilura Elliot, 1871^ 

1792. Felis bengalensis. Kgh. Animal Kingdom, p. 151. Southern Bengal. 
1837. Felis chinensis. Gray. Mag. Nat. Hist., vol. 1, p. 577. China. Fixed 

as Canton (Allen, 1938; Ellerman and Morrison-Scott, 1961 and 

1862. Felis widata. Radde. Reisen Siiden OstSibirien, vol. 1, p. 106. Amur, 

60 km below mouth of 2^ya. Nee Felis undata Desmarest, 1816. 

1871. Felis euptilura. Elliot. Proc. Zool. Soc. London, Dec. 1871, p. 761. 
Substitute for undata Radde. 

251 1872. Felis microtis. Milne-Edwards. Rech. Hist. Nat. Mamm., p. 221. 
Near Peking. 

1872. Felis decolorata. Milne- Edwards. Ibid., p. 223. Near Peking. 
1904. Oncoides bengalensis Kerr a. raddei. Trouessart. Catal. Mamm. 

Quinq., Suppl., p. 271. Replacement for undata Radde, 1862. 
1922. Felis manchurica. Mori. Лил. Mag. Nat. Hist., vol. 10, p. 609. Near 

1929. Felis viverrina mandjurica. Baikov. Okhotnik, vol. 4, p. 16. 

Northeastern China (Manchuria). (V.H.) 


Body spotted throughout but spots on back may be faint; color of spots rusty. 
White field present on back of ear. Two light-colored stripes run from inner 
comer of eyes onto forehead. Skull elongated; plate in inteфterygoid region 
extended posteriorly (Fig. 158**); and inteфterygoid vacuity very narrow. 

*In the most recent period, as a result of assigning the name bengalensis to this species, 
it is sometimes refeired to as Bengal cat. Such a modification of the Russian name, which 
came into common usage long ago (Satunin, 1914; Ognev, 1935; Novikov, 1956; Stroganov, 
1962; and others) is unnecessary and only leads to confusion. 

^The species name euptilura is assigned with some reservation. It is possible that the 
name microtis has some completely formal advantage over it. According to some data the 
work of Milne-Edwards published between 1868 and 1874 should be regarded as belonging 
to 1871, especially the eariier sections (Trouessart, 1905; April — Sowerby, 1923), but Elliot's 
work was published in December, 1871.* According to Ellerman and Morrison-Scott (1951 
and 1966), however, the name microtis is dated 1872. Finally, sometimes the names assigned 
by Milne-Edwards in the above work are assigned simply to 1874. In any case, continuation 
of the name euptilura is desirable as a nomen conservandum. There is sufficient justification 
for concluding that the form chinensis belongs to another species (bengalensis; Heptner, 
1971). If, however, it were proven that it was conspecific with the Amur form, it is clear 
that the name chinensis Gray, 1837 should have distinct priority as a species name. 

*1971 in Russian original — Sci. Ed. 

**Misprinted 156 in Russian original — Sci. Ed. 











Fig. 156. Amur cat, Felis {Felis) euptilura (yearling female. "Kedrovaya Pad' 
" preserve near Vladivostok. October 27, 1%5. Photograph by A.G. Pankrat'ev. 

Second premolar (first in row) usually present; antero-inner cusp of upper 
carnassial tooth weakly developed and without prominent cusps. (V.H.) 


Small animal, roughly the size of Caucasian forest cat [Fells sllvatlcus 

In general build and appearance this species is similar to the Caucasian 
forest cat but stands slightly taller on its legs, though not as tall as the jungle 
cat, and is lighter (Fig. 156"'). Tail somewhat shorter than one-half of body 
length, but usually more than double the length of hind limb. Head small, 
fairly narrow at the zygoma, and hence facial region pointed and somewhat 
extended. Ears quite large, close-set, and project prominently from pelage, 
even in winter; tip of ear somewhat blunt and devoid of tuft of long hair. 
Hair on cheeks not long (side- whiskers absent). Eyes small with a vertical 
pupil. Pelage throughout body of uniform length; tail furry and appears 
quite thick. 

' 'Pelage similar to that of European forest cat. On the back an average 
of 7,000 hair occur per cm^ and on the abdomen, 1,950. There are 30 

*Misprinted 157 in Russian original — Sci. Ed. 


252 Fig. 157. Head of an old male Amur cat, Felts (Felis) euptilura. ' 'Kedrovaya Pad ' ' ' 

preserve near Vladivostok. October 3, 1965. Photograph by A.G. Pankrat'ev. 

underfur for every top hair. Pelage on back consists of six categories of hair, 
and five on the abdomen (category IV top hair absent). Average length of 
guard and top hair on back 47, 49, 47, 39, and 35 mm, and their thickness 
108, 106, 75, 66, and 34 microns; corresponding values for hair on the 
abdomen 45, 43, 37, and 31 mm, and 60, 56, 40, and 41 microns; length 
and thickness of underfur on the back 34 mm and 18 microns, and on the 
abdomen 28 mm and 16 microns. Thus there is very little difference in the 
length of hair on the upper and lower surfaces of the body. The shortness 
and thickness of guard hair and the absence of one category of top hair on 
the abdomen are characteristic features" (B.F. Tserevitinov). 

Typical coloration spotty but extremely variable. In extreme cases the 
entire body of one animal was covered with spots, while spottedness was 
almost absent in another, standing out prominently only in a small area. The 
color difference between extreme variants is so great that it has provided 


a basis for some zoologists to propose the existence of two types of small 
forest cats — one in the Ussuri territory and the other in northeastern China 
(Maak, 1861; Baikov, 1915 and 1929). 

The spotted type of coloration, when fully developed, is as follows: 
Background color on back and upper part of flanks in winter coat usually 
variable — dirty grayish-pale-yellow or dull grayish-brown with a slight rusty 
or reddish hue. Color gradually fades downward on flanks, becoming whitish 
ocherous. Abdomen, groins, and inner sides of legs very light-colored, 
ocherous or dirty white, or white with a light ocherous dusting. Chin, throat, 
underside of neck, and chest have a white background. 

Throughout the body, extremities, along the neck, and in part along 
the head, occur small (1.0 to 2.0 cm) complete spots of circular or slightly 
elongated shape; the latter are particularly evident on the back. They have 
in part shaф outlines, and in part their margins are blurred and indistinct. 
They are either riLSty-ocherous or rusty-cinnamon, light and bright, or duller 
and fuller. They may be very close-set; sometimes the main background 
color occupies almost the same area as the total area of the spots. The spots 
are denser and their total area relatively greater in this species than in other 
Soviet species of small cats. 

Three or four narrow stripes extend along the spine consisting of 
elongated, narrow, 8Ьаф, and richly colored small spots. Sometimes their 
outlines are so indistinct that a generally brown, indistinct, broad belt is 
formed. Spots and longitudinal rows of shaфly outlined spots stand out 
prominently only at places, mainly on the shoulders and sacrum. Spots on 
the abdomen and chest are more rounded than on the back, bright, and rusty 
or rusty-brown in color. Down the throat and neck, the spots are bright rusty 
or a much darker rusty tone. They are usually disposed in such a way as to 
form three to five transverse rows, but often fuse into continuous transverse 
i53 stripes. Rows of spots, three to five, fusing or almost fusing into stripes, 
sometimes occur across the upper part of the forelegs. Spots on the 
extremities become increasingly smaller and extend to the tarsal and carpal 
region. Vertical stripes or vertical rows of spots in a pattern do not occur. 

The tail is dark gray, turning somewhat lighter in color toward the lower 
part, and is either monochromatic or with fairly distinct dark (blackish- 
gray) transverse broad stripes or bands. Tip pure black or dark gray. 
Throughout the tail an extremely faint ocherous dusting is sometimes present 
and the stripes thus may have an admixture of rusty tones. As many as seven 
stripes may occur on the tail. 

Coloration of head also complex (Fig. 157*). Upper lip, area around 
eyes, as well as lower lip white, but riLSt-colored spots occur in the region 

*Misprinted 158 in Russian original — Sci. Ed. 


255 Fig. 158. Skull of Amur cat, Felis (Felts) euptilura. No. S 14197, collection of 

the Zoological Museum, Moscow University. Cape Gamov near Vladivostok. 
February 12, 1927. Sketch by N.N. Kondakov. 

of the whiskers. Upper part of nose grayish with a rusty dusting. From the 
nose through the forehead to top of head there occurs a broad, dark, reddish- 


cinnamon stripe, set off on both sides by two distinctly prominent narrow, 
light-colored (white) fields. This particular marking commences at the inner 
corner of the eye and extends through the forehead. Beyond, the stripe and 
white fields diminish and disappear in the general dark color on top of the 
head. The light4:olored fields are bordered by a narrow dark field, somewhat 
darker than the space between the eye and the ear and the entire upper part 
of the head. 

Down the cheeks and behind the eyes a light-colored field occurs, along 
which two bright stripes of different intensities of reddish, rusty, or light 
brown color extend posteriorly from the region of the eye. This particular 
marking merges into the pattern on the throat and the lower side of the neck, 
or touches it in some manner. On the dark-colored sinciput, occiput, and 
upward on the neck, sometimes almost up to the shoulder, four narrow brown 
or rusty-brown fields occur. The back of the ear is, for the most part, white 
or almost white; just below the tip it is blackish-brown; the tip itself is 
light rusty in color. 

The second type of coloration, when fully developed, is as follows. 
Back and upper flanks indistinct, dirty, brownish gray, with an admixture 
of silver at places due to the presence of many light -colored top hairs. Color 
richer and darker on the back, and somewhat lighter on the flanks, especially 
ventrally. Spots not visible against this main background. Body essentially 
monochromatic, brownish-gray. Spots faintly discernible along undersides 
of flanks but distinct on abdomen, where their brown shade stands out 
prominently against the ocherous white field. Spots indistinct on legs. Spots 
distinct against general background of light-colored body; nevertheless they 
are rather vague, diffused, and rusty in tone. Some spots on abdomen and 
thighs have a small light-colored center. 

Tail dark, dirty gray, with darker transverse spots. Reddish-colored spots 
are absent throughout the body in this color pattern; instead, spots are 
brownish-rusty on the throat, down the neck, and on the groin. The head 
is a dark or coffee -brown color and the stripes along the sinciput, occiput, 
and up the neck, dark brown or even blackish-brown. Very distinct light- 
colored fields extend from the eyes to the forehead. 

The two extreme types of color patterns described above are often 
blended in various ways. The main background may vary from dark gray 
or dark brownish-gray to almost dirty black along the spine (up to the 
"belt"), to light reddish-ocher and pale yellowish-ocher. Spots may be 
close-set through the back, flanks, and down the body, or sparse, or not 
distinguishable, or only barely discernible on the spine; sometimes spots 
are more "blotches" than spots. In dark individuals spots may be totally 
absent down the middle of the back but quite distinct on the flanks, or absent 
throughout the body except for the abdomen. Sometimes spotting is represent- 


ed by faint shading. Rarely, spots on the thighs and abdomen will have 
a small, indistinctly contoured, lighter-colored area in the center. Spots may 
vary from vivid red to dark brown, without entirely "reddish" [intermediate 
tones] (even on the abdomen) and dirty black. Stripes on the occiput and 
neck may be distinct, black, and long, reaching up to the shoulder, or almost 
imperceptible. Corresponding to the general color tone and inteasity of color 
of the coat, the color pattern on the upper part of the head also varies. Stripes 
on the cheeks may be bright reddish or rusty, or dark and brown, but 
invariably exhibit a distinct rusty tone. Spots and transverse bands on the 
throat vary from bright ocherous-rusty to dark brown with a rusty hue. The 
254 tail likewise varies in color, from an almost monochromatic gray in dark- 
colored animals to gray with faint spots, even though the body spots in 
some animals may be very vague. 

Judging from available, albeit meager data, different types of variegated 
color patterns are found in the population, while dark, monochromatic 
animals are quite rare. According to some authors (A.G. Pankrat'ev) the 
coat may fade significantly after winter. However, both spotted and mono- 
chromatic cats are encountered in early winter. 

On the whole, color variation in the Amur cat is greater than that of 
any other species of Soviet cat, except lynx. However, in all color types 
the light-colored stripes extending from the comer of the eye and the reddish 
hue of the pattern on the throat and down the neck remain unchanged. The 
rusty shade in the fields along the cheeks is also constant, as is the color 
of the back of the ear.^ 

Not much is known about the summer coat. It is evidently characterized 
by the presence of sharply visible spots, but the overall range of variation 
characteristic of the winter coat is also seen in summer. 

Sexual dimoфhism is absent and age-related changes have not been 
described for Soviet forms. In any case, by the first winter young kittens 
have developed an adult coat. Geographic color variation throughout the 
species ' range under consideration here is not adequately known but seems 
to be insignificant. 

The skull is notably elongated, with an extended braincase and close- 
set zygomatic arches, and is low. Upper line of profile bulges slightly and 
uniformly without prominent elevations in the interorbital region. Frontal 
region small, narrow, gently bulging, and median longitudinal groove absent. 
Supraorbital processes well developed, long, usually approaching postorbital 

^hese descriptions of coloration of Amur cat vary greatly from those given in Soviet 
literature, including even those given for purposes of identification. In the latter a rich and 
vivid reddish spottiness throughout the body is invariably emphasized as the main 
characteristic. The present description is based on data from the Zoological Museum, Moscow 
Univereity, but mainly on new data furni.shed by A.G. Pankrat'ev (Vladivostok). 


processes of the jugal, but sometimes (more often in old animals) fused, 
in which case orbit is closed posteriorly. 

Sagittal crest in posterior part of parietals distinct but low; lambdoidal 
crests well developed and more complete than sagittal. Rostral part of skull 
relatively short and broad. Orbits oval, significantly extended backward, 
and their ventro-anterior region greatly thickened. Maximum diameter of 
infraorbital foramina wider than space between these openings and edge 
of orbits. Nasals narrow in posterior half but extended and broad in anterior 
part. Hard palate narrow and long — its posterior margin lies in inteфterygoid 
vacuity beyond the line joining posterior surface of molars, at a distance 
more than one-half width of interpterygoid vacuity (see Fig. 158). 

Interpterygoid vacuity narrow. Choanal openings small; diameter only 
slightly more than height. Tympanic bulla large, greatly inflated, and long; 
anterior margin lies close to postglenoid process but does not reach its level. 
Ectotympanic chamber small but its boundary with entotympanic chamber 
not visible externally. Distance between inner margins of tympanic bullae 
equal, or almost equal, to width of interpterygoid vacuity. 

Canines thin, shaф, and moderate in length. Second (first in row) upper 
premolar usually present but may be shed during lifetime and even its cavity 
filled in most adults, especially older animals. Both teeth absent in 30 to 
40% of animals examined (A.G. Pankrat'ev). Antero-inner cusp of upper 
carnassial negligible, or weakly developed, and without shaф edge. 

Sexual dimorphism in skull absent apart from somewhat smaller size 
of female skull. Skull of young animals not as long but with larger cranium. 
Even in the first winter, differences between the skull of young animals 
and that of an aduh diminish notably. Geographic variation of skull mainly 
concerns overall size, which can be rather significant. 

For animal measurements see "Geographic Variation". (V.H.) 

255 Systematic Position 

The Amur forest cat was initially (Radde, 1862) placed under Felis undata 
Desm., a species of the Malay Archipelago. Later it was clarified that the 
Amur forest cat is not related to this species and hence it was (by means 
of a substitute name) treated as a distinct new species, Felis euptUura (Elliot, 
1871). In the various nomenclatural schemes adopted in Soviet and foreign 
literature, the Amur forest cat was treated as a distinct species of forest cat 
found in the Amur-Ussuri territory and northeastem and eastern China until 
the early 1900's, when it was suggested that the entire group of forms in 

256 the species euptilura be regarded simply as a subspecies of the south Asian 
species, F. bengalensis Kerr, 1792 (Trouessart, 1904). This suggestion was 
accepted by several prominent zoologists (Pocock, 1939; Ellerman and 


Morrison-Scott, 1951; and others), albeit rather tardily. In foreign literature 
this was generally accepted. Soviet zoologists, on the other hand, clung to 
the independence of the species F. euptilura (Ognev, 1935; Bobrinskii et 
al., 1944; Novikov, 1956); however quite recently, a few have now begun 
to favor the view that the Amur cat should be assigned to the species 
bengalensis (Bobrinskii, Kuznetsov, and Kuzyakin,1965). 

A special study of this problem (Heptner, 1971) has revealed that the 
Amur cat is more correctly isolated as an independent species, differing 
significantly from F. bengalensis of southern Asia. The Amur cat is 
characterized by a number of peculiarities, which distinguish it not only 
quantitatively but also qualitatively. Compared with F. euptilura, the skull 
of F. bengalensis has a relatively larger cranium, the lyriform figure on it 
is larger, and the lines forming it, in fully grown animals, converge not 
in the region of the frontals or the coronal suture, but farther behind, roughly 
at the middle of the parietals. The sagittal crest is not developed and appears 
only at the extreme posterior end. Compared with the skull of the Amur 
cat, therefore, the skull of F. bengalensis is slightly more juvenile — the 
skull of adults resembles that of younger Amur animals. In this respect, as 
shown above (see characters of the family) the skull of F. bengalensis does 
exhibit some "progressive" features. In these it is similar to the skull of 
cats of the F. silvestris type. The skull of the Amur cat exhibits structural 
features which are transitional between those of F. bengalensis and F. 
viverrina and, in some respects, are closer to the skull of the latter species. 
However, it is significantly smaller and the crests are less developed. 

Of greater significance are the color differences between Amur cat and 
F. bengalensis. Coloration and spottedness are completely different in the 
Bengal cat. Spots in the latter are invariably shaф against a very light- 
colored background and readily differentiated in form and size. Small spots 
occur with very large spots, mostly irregular in form, but often elongated, 
especially on the spine and close to it. Tail coloration also differs. In the 
Amur cat some individuals are almost monochromatic; dark or very faintly 
spotted patterns are normal. If, however, the spots are distinct they are 
numerous, identical, small, densely set, and do not tend toward elongation. 
This is the more prevalent type of color pattern (see "Description"). This 
type of spotted pattern (leaving aside the relation of spots and background) 
is similar to that in the steppe cat (F. silvestris caudatd). In general, however, 
the coloration, in particular in the sharply spotted type found in the Amur 
cat, quite closely resembles the coloration of F. viverrina!^ Color 
characteristics, of vital importance among cats, reveal a greater affinity with 

^See sketches off. bengalensis and F. viverrina presented by R.I. Pocock (1939) and 
coloration descriptions of species F. bengalensis and F. euptilura given by V.G. Heptner (1971). 


F. viverrina than in skull stmcture. On the whole, while belonging to the 
same group, ''Pno?iailunis,'' as do F. viverrina, F. bengalensis, and F. 
rubigitwsa (possibly also F. planiceps), F. euptilura occupies an intermediate 
position between F. viverrina and F. bengalensis. 

Moreover, the delimitation of this group, which at times is considered 
a subgenus of genus Felis and at times an independent genus, is not well 
defined; neither is its compactness. Thus some workers break this "genus" 
into several separate subgenera distinct from genus Felis — viverrina 
(Zibethailurus) and planiceps (Ictailurus). Moreover, the entire group 
"Prionailurus" exhibits a very close relationship with the conventional group 
Felis s. str. and, as mentioned earlier, its further classification into species 
is difficult to substantiate. Hence, here, F. euptilura is placed in genus Felis 
s. 1. and subgenus Felis. 

In skull stmcture (see characters of the family and genera Panthera and 
Felis), F. euptilura occupies, among Soviet species of the genus, the position 
of least specialized or one of the least specialized forms. This species stands 
257 at the beginning of the series which leads to the more progressive conditioas 
culminating in development of the "feline" type in genus Felis s. 1. — 
the manul; it is closest to F. chaus (jungle cat). 

The proximity of Amur cat to species oi Felis s. str. and the incorrectness 
of separating it as an independent subgenus, or even a genus, are supported 
by the fact that the Amur cat hybridizes with the domestic cat in natural 
conditions. Crossbreeding often takes place in forest villages in the Ussuri, 
the male partner being the wild cat. In a backcrossing of hybrid males and 
females with domestic males and females, progeny have been obtained which 
are fertile in both combinations. Furthermore the hybrid generation is fertile 
among its own members (A.G. Pankrat'ev). 

For body measurements, see "Geographic Variation". (V.H.) 

Geographic Distribution 

The Amur cat inhabits forested regions of the Amur- Ussuri territory and 
northeastern and eastern China. 

Geographic Range in the Soviet Union 

The range in the Soviet Union (Fig. 159) represents the northeastern 
boundary of the range of the species. It encompasses the southern part of 
the Far East and associated forests of the so-called Manchurian type. The 
range is intermpted within the Soviet Union but the two sections merge 
outside the boundaries of our country. 

In the west the range boundary, commencing at Blagoveshchensk on 
the Amur, runs toward the middle course of the Tom', the left tributary of 


Fig. 159. Distribution of Amur cat, Felts (Felis) euptilura, in the Soviet Union 

(scale in km). 

1 — northern boundaiy of range according to data up to the early 1930's; 2 — same, 
up to 1968; 3 — discontinuous (diffuse) distribution; 4 — dense distribution and 
region of highest population density in 1%8; 5 — possible habitation up to Gorin 
mouth in the past. V.G. Heptner (distribution and population density in 1968 
according to data of A.G. Pankrat'ev and G.F. Bromlei). 

the lower Zeya, roughly toward 51°17' N. lat. (northernmost point of 
258 occurrence). Turning east and slightly south it then intersects the Turan range 
and the middle course of the Bureya, runs toward the upper Mutana, and 
descending from there to 49° N. lat., turns toward the upper Bidzhan. From 
there it descends southward and enters China. 

On the right bank of the Amur, the northern boundary of the range is 
formed by the Amur in the stretch from the state boundary and the mouth 


of the Ussuri in the west to the mouth of the Sungari in the east. From 
this point (50°06' N. lat.) the boundary runs southeast and south along the 
sources of the Monoma, Anyuy, and Югог, descending to the Mukhen' River 
around 48°55' N. lat. From there it runs eastward, intersects the upper Khor, 
and through the Murav'ev-Amur pass to the Samarga River and the sea 
of Japan at 47°20' N. lat. In the expanse between Ussuri and the ocean, 
the Amur cat is confined to nut pine- broad-leaved, and mixed forests, and 
does not ascend beyond 500 to 550 m above sea level. In other words this 
species is absent in the higher altitudes of the Sikhote-Alin covered by 
Okhotsk taiga. It is possible that, in the north, it formerly reached the mouth 
of the Gorin (50°45' N. lat.). This species breeds on Russkii Island at 

The boundary outlined above, although schematic (range in Ussuri 
territory around Sikhote-Alin), summarizes data available on the distribution 
of Amur cats from the time of the first Russian explorers of the Far East 
(1850's) to the eariy 1930's (Ognev, 1935; Zubarovski, 1939). 

The 1968 boundary was probably of the following form. Commencing 
in the west slightly east of Blagoveshchensk on the Amur, it ran southeast 
along the river at distances from it varying from 30 to 50 km, to 131° E. 
long. There it sharply deviated from the river, turning initially straight 
eastward and later southeast, and, intersecting the Amur, entered China. 
This arcuate boundary covered the entire Bol'shoi Bira basin except the 
upper portions of its left tributaries, and crossed the Amur at some 40 to 
50 km below the mouth of the Bol'shoi Bira. Within the Soviet Union, 
Amur cats were not found at the time of the survey farther down along 
the left bank of the Amur River. 

The boundary again re-entered the Soviet Union at the mouth of the 
Ussuri but Amur cats were absent in the immediate proximity of IChabarovsk. 
On the right bank of Amur, the boundary extended to the mouth of the 
Sungari. There it turned slightly eastward into the mountains, and then tumed 
sharply southward along the western slope of the Sikhote-Alin through the 
upper Anyuy, Khor, Bikin, and Iman, without embracing, however, their 
sources and uppermost reaches. The boundary extended in this direction 
south to 45° or 45°15' N. lat. The boundary ran here for a distance of 120 
to 150 km from the Ussuri. At this point in the south it intersected the main 
axis of the mountains and turned immediately northeast, running along the 
eastern slope of the Sikhote-Alin. It continued north in this direction to 
the mouth of the Botcha (Grossevichi) at 48° N. lat., where it reached the 
sea. This part of the range is in the form of a very narrow strip at its widest 
point in the south (about 60 km wide slightly north of Temeya Bay). Thus the 
extensive, higher parts of the Sikhote-Alin, covered by Okhotsk type taiga 
or devoid of forests, did not fall within the range. Farther south the range 


covered the entire territory of the Ussuri krai (from data of A.G. Pankrat'ev 
and G.F. Bromlei). 

Boundary differences between the 1930's and end of the 1960's are 
primarily the result of precise demarcation in recent decades (Sikhote-Alin, 
left bank of Amur). However, some reduction in the range has also been 
brought about by the extinction of the animal in the north where its population 
has always been sparse (see subdivision "Population" under the section 
"Biology" below), from changes in natural conditions (clearing of forests 

259 Fig. 160. Di-stribution (scale in km) of Amur forest cat, Felis (Felis) euptilura Ell., 

and Bengal cat, Felis (Felis) bengalensis Kerr. Lower reaches of Yangtze — 
southernmost point from which Amur cat is known (Nanking; Heptner, 1971). 
Southern boundary of range of Amur cat calculated as SO^N. lat. V.G. Heptner. 


and shrubs). Adverse effects in one part of the range are naturally reflected 
in other parts. Evidently boundary variations, although not directly proved, 
also arise from natural climatic and landscape (forests of Manchurian type) 
limits to the range. Existing under such conditions the species must have 
become sensitive not only to long-term climatic changes (they could have 
remained adequate for over a hundred years) but also to temporary 

259 Geographic Range outside the Soviet Union 

How large the size of the range was outside the Soviet Union (Fig. 160) 
(related to the question of forms) cannot be stated with certitude since the 
species limits are not yet clear (see "Systematics" and "Geographic 
Variation"). In any case, it includes northeastern China (evidently only 
the eastern and northeastern parts), Korean Peniasula, the islands of Cheju Do 
and Saisyu (Tsashima and Quelpart), and eastern China, at least to the lower 
course of the Yangtze Kiang (Nanking ^^). It is possible that this cat occurs 
even farther south but it is not known precisely to what extent and where 
the boundary lay in the past in relation to that of F. bengalensis, nor whether 
these two species are (were) encountered together in southern China. In any 
case, F. b. bengalensis has been recorded from Yunnan; the form described 
under the name F. b. chinensis from Fukien, Hunan, Shansi, Hopei, and 
Szechwan, a small cat with some color features oi bengalensis and euptihira, 
judging from descriptions, pertains to bengalensis (Allen, 1938). The 
distribution of the species in western China is also not clearly known. True 
F. bengalensis, with wholly typical features, lives in the Democratic Republic 
of Vietnam. 

260 Cats assigned to the species bengalensis, in addition to the foregoing regions, 
also inhabit Gansu and Szechwan, Kam, Taiwan (Formosa) and Hainan, 
Indochina and Malacca, the islands of Sumatra, Java, Bali, Kalimantan 
(Borneo), Palawan, Calamian, Panay, Negros, and Cebu in the Philippines, 
Burma, Assam, Sikkim, Bhutan, Nepal, Kumaon, Kashmir, Punjab, the Indian 
Peninsula almost up to the southern extremity (Palni hills at 10° N. lat.), 
and southern Baluchistan. This species is absent in Sri Lanka. References 
to its occurrence in Tibet are erroneous or applicable only to Kam. 

Although the extent of distribution of the species euptilura, in the sense 
accepted here, is not wholly clear, it is obvious that true F. bengalensis lives 
in the Democratic Republic of Vietnam and evidently also in southeastern 

^•^Specimen in collection of the Zoological Museum, Humboldt University (Berlin, 
German Democratic Republic) (V.G. Heptner). 


tropical and subtropical China. Thus the range of F. euptilura in relation 
to that of F. bengalemis is allopatric; however the overlapping of their ranges 
in the south cannot be ruled out. It is interesting that the range of F. euptilura 
coincides with the region of occurrence of Manchurian-Chinese relicts. The 
species itself should probably be considered a member of this relict group. 
It has been suggested (Pocock, 1939) that F. bengalemis, i.e., all forms 
of southern Asia from Kashmir to southern Baluchistan, Indochina, the Malay 
Archipelago, and southern China, and the group euptilura, regarded here 
as an independent species, evolved (like tiger) in the north and, later, having 
colonized southward in east Asia, extended still farther southward. As can 
be seen from the above account such a viewpoint is difflcult to accept. If 
F. bengalemis s. str. and F. euptilura have common roots, thenF. bengalensis 
should also be very old in origin (typical of south Asian species), like several 
other forms of the eastern (Indo-Malayan) region, i.e., F. rubiginosa and 
F. viverrina, which are close to it, as well as F. nebulosa, F. marmorata, 
F. temminckii, F. chaus, and others. A reverse situation is possible: a distinct 
colonization of some progenitors of south Asian cats into the north for F. 
euptilura. (V.H.) 

Geographic Variation 

The species range of the Amur cat, in the accepted interpretation of the scope 
of the species, is not very large. Nevertheless it extends fairiy far south 
where environmental conditions significantly differ from those prevailing in 
the Amur-Ussuri territory. Its geographic variation has not been adequately 
analyzed, apart from the normally occurring range of individual variation. 
At present it is still difficult to assess positively which among the subspecies 
assigned traditionally to bengalensis s. 1. actually belong to this species, 
and which to euptilura. This pertains in particular to the form chinensis. 

Only one subspecies occurs within the Soviet Union. 

Amur forest cat, F. (F.) e. euptilura Elliot, 1871 (syn. raddei, 
manchurica, manjurica; the names undata, bengalensis, chinensis, and 
microtis were once applied to this form). 

The "Description" given above pertains to this form and has been 
developed on material from the Ussuri territory. 

Size large, representing the largest form of the species. 

Body length of males 60 to 85 cm, and of females 61 to 77 cm; tail 
length 23.5 to 44.0 cm and length of ears 4.2 to 5.0 cm. Height at shoulder, 
31 to 36 cm. 

Greatest length of skull in males 96.7 to 115.3 mm, and of females 88.2 
to 102.0 mm; condylobasal length of skull in males 89.5 to 105.0 mm, and 
in females 82.3 to 92.8 mm; zygomatic width in males 60.7 to 74.5, and 


in females 57.0 to 67.2 mm; interorbitai width in males 14.4 to 18.8 mm, 
and in females 13.1 to 16.3 mm; postorbital width in males 27.0 to 28.8, 
and in females 26.0 to 28.0 mm; width between infraorbital foramina in 
males 27.5 to 29.0 mm, and in females 25.7 to 27.3 mm; width above canines 
261 in males 24.1 to 28.6 mm, and in females 24.1 to 26.0 mm; and length 
of upper tooth row (with canine) in males 31.0 to 34.5 mm, and in females 
27.8 to 31.0 mm. 

Weight varies from 2.5 to 7.07 kg; one animal weighed 8.2 kg and 
another 9.914 kg (exceptionally obese male).^^ 

Much bigger, gigantic individuals are extreme rarities in the Soviet 
Union. Thus, at Birobidzhan a male was caught (December, 1935) with a 
body length (from nose to anus) of 107 cm and tail 44 cm, and another male 
north of Vladivostok (Anuchinsk region; February, 1936) with a body length 
of 103 cm and tail 42 cm. These animals were measured "in the flesh". 
The skins of some cats correspond in size to those of small lynx (about 100 
cm without tail; Zubarovskii, 1959). Such specimens fall so much out of 
the normal range that they cannot even be regarded as extreme variants. 
It is possible that they represent heterotic hybrids of Amur with domestic 
cat (see "Systematics" above). 

Measurements of forest cat populations of the southernmost part of the 
Ussuri region are presented in Tables 5 and 6P 

The extremely high variability in weight of the cats is due not only to 
sex-related and age-related differences, but seasonal changes. The maximum 
weight of old males occurs at the beginning of winter. By spring it drops 
greatly, although sometimes the animals are well fed even in January. Much 
depends on climatic and other conditions in a given year; cats become very 
obese when small rodents are plentiful; otherwise, they become extremely 
emaciated even in January and Febmary (A.G. Pankrat'ev). "Exceptionally 
obese" males and females were caught on October 25, 1946 and January 
2, 1945 with a body length of 66 and 62 cm, weighing 5,700 and 4,319 
g (G.F. Bromlei).^-^ 

Weight of heart of adult and subadult males (10) 8.3 to 37.0 g (M 21.04) 
and cardiac index 3.49 to 7.89 (M 5.35); corresponding values for adult and 

^^Suramarized data include those published by N.A. Baikov (1934), S.I. Ognev (1935), 
M.I. Zubarovskii (1939), S.U. Stroganov (1962), material furnished by N.K. Vereshchagin 
(collection of the Zoological Museum, Academy of Sciences) and A.G. Panki^at'ev. and collection 
of the Zoological Museum, Moscow University; in all, some 50 specimens are represented. 

^^ables include specimens of the Zoological Museum, Moscow University, data 
furnished by G.F. Bromlei (Sudzukhin preserve), and material of A.G. Pankrat'ev (vicinity 
of "Kedrovaya Pad' "' preserve, southwest of Vladivostok). 

^•'It changes with the nutritional plane, and also affects the biologically important 
character of weight load on the sole (see "Biology"). 

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subadult females (10) 10.6 to 27.3 g (M 18.16) and cardiac index 4.54 to 
6.23 (M 5.18). 

Length of intestines in adult and subadult males (11) 120.0 to 189.0 
cm (M 159.5); ratio of same to body length 1 : 2.30 to 1 : 3.0 (M 1 : 2.57); 
corresponding values in adult and subaduU females (10) 132.0 to 21 1.0 cm 
(M 149.6) and 1:2.36 to 1:2.70 (M 1:2.55) (A.G. Pankrat'ev). 

F. euptUura is found in the Pri-Amur and Ussuri territory. 

Outside the Soviet Union these animals occur in northeastern China 
(Manchuria) and the Korean Peninsula. 
263 In the accepted inteфretation of species euptUura, and in accordance 

with the earlier nomenclatural discussion (see pp. 335 to 337), the nominal 
form inhabits the Soviet Union. In any case, labeling them ''Fells euptilura 
microtis,''' as has been the practice, or placed in form chinemis, as suggested 
recently (Bobrinskii et al., 1965) should not be done (Heptner, 1971). 

Relations between the Soviet form and several geographically adjacent 
forms are not clearly understood. Evidently this same large form has been 
found in northeastern China also (Baikov, 1929) and a similar, but much 
smaller, form seen in the south. ^"^ This may be called microtis M.-E. (type 
loc. Peking). Sometimes it is labeled chinensis Gray (type loc. Canton), 
which is not correct. The name manchurica Mori is evidently a synonym 
of euptilura. 

Either 10 or 11 forms are usually included in species bengalensis 
outside the Soviet Union (Chasen, 1940; Ellerman and Morrison-Scott, 
1966; Weigel, 1961). This number is evidently an exaggeration. Of these 
forms chinensis (see above p. 342 and Heptner, 1971) possibly belongs 
to species euptilura. It occurs from Fukien in the east to eastern Tibet and 
Szechwan in the west, Mongolia (Inner Mongolia — V.H.), and Taiwan in 
the north [sic] (V.H.). 


Population. The range of the Amur cat is associated with Manchurian type 
broad-leaved forests. In the Bureya basin and the Zeya and its tributary, 
the Tom' this cat is very rare, and even rarer in Birobidzhan (almost extinct). 
On the Zeya-Bureinsk plain ("Amur forest-steppe") and in Birobidzhan, 

^''Measurements reported by Shou (1958)* evidently pertain mainly to young animals: 
body length (seven animals) 46.0 to 55.5 cm; weight (five animals) 1,750 to 2,875 g; and 
total skull length (four animals) 87.1 to 98.1 mm. In addition to the foregoing, mention is 
made of one animal with a body length of 66.0 cm and another weighing 7,200 g (female 
with a body length of 55.0 cm) (V.H.). 

*Not in Literature Cited — Sci. Ed. 


however, this cat was common before the war. The reasons for its population 
reduction are the cultivation of virgin lands, burning and felling of forests 
and thickets which mainly serve as shelters for the animals, and the plethora 
of hunters who invaded the remaining small areas of habitat. On being chased 
by a dog, an Amur cat invariably escapes into a tree; here it is easily killed 
and usually thrown away. This species avoids large dense forests (A.G. 
Pankrat'ev). In the basin of the lower Ussuri it has become fairly rare in 
recent years. It is still quite common in the southern half of the Ussuri territory 
(Pos'etsk, Vladivostok, Shkotovsk, Suchansk, and other regions). In each 
of these regions in the 1920's some 100 to 150 cats were caught annually. 
The maximum density in years of high population in the extreme south of 
the Ussuri territory, in "Kedrovaya Pad' " preserve south of the Khasansk 
region, was two to five animals per 1,000 hectares (A.G. Pankrat'ev). 

Habitat}^ The Amur cat, like most other cats, is not adapted to living 
in very snowy regions, i.e., in areas where the snow cover over a 10-day 
period exceeds 20 to 30 cm, or snowfall is protracted. However, this cat 
finds optimum conditions of existence in sections where the snow cover 
is less than 10 cm deep (Formozov, 1946). With a very high weight load 
on the sole of the foot, equalling 188 g per cm^ (G.F. Bromlei),^^ the animal 
sinks in deeply in loose snow, and hence avoids snow covers of greater 
depths. Confronted with a snow cover the cat usually moves along crusted 
portions on streams and brooks, trails of other animals, roads, and rocky 
sections, from which the snow has been blown off by the wind, or under 
264 fallen trees and places cleared of snow by marine surf. Deep snow cover 
deprives this animal of its chief food, i.e., rodents, since the predator cannot 
catch them under snow. 

A second requisite for the occurrence of this cat in a given region is 
a fairly abundant supply of mouse-like rodents, Manchurian hares, and 
pheasants, which it usually hunts. The selection of habitat by this animal 
is related to these requirements. Thus in the southern Primor'e and Ussuri 
basin it inhabits sparse Manchurian type broad-leaved forests where these 
dietary items are more common. The predator occurs less often in these 
regions in nut pine-broad-leaved forests. It does not ascend mountaias beyond 
500 m above sea level. It is frequently encountered on creeks draining into 
river valleys, among thickets of rose willow or sedge, where h hunts for 
hare. In winter this cat is sometimes confined close to unfrozen springs from 
which it can catch fish (Zubarovskii, 1939). It rarely appears on the western 

^•''Habitats of the Amur cat are also characterized by photographs of the biotopes of 
Amur leopard and Amur tiger; the three species are encountered together almost everywhere. 

^^According to A.G. Pankrat'ev, in adult and subadult males (10 animals), depending 
on tlieir obesity, the weight load per cm^ of sole surface was 48 to 115 g, and that in females 
(10 animals) 50 to 87 g. 


Fig. 161. Habitat of the Aniur cat in tlie lower Ta-Chingou River. Suputin preserve, 
Ussuri territory. August, 1958. Photograph by V.E. Prisyazhnyuk. 

slopes of the Sikhote-Alin (N.V. Rakov) because climatic conditions 
prevailing there are not favorable. Winter in that region extends from mid- 
November to March and is severe and frosty. Even at the end of November 
the atmospheric temperature may drop to -35° С Winter sets in quickly 
and snowfall is continuous throughout the season in Sikhote-Alin. Thus few 
Manchurian animals are seen in this region during the winter. 

Close to the Sea of Japan, on the eastern slope of Sikhote-Alin, the 
Amur cat lives mostly along the sea coast. A stretch 5.0 to 6.0 km wide 
along the sea, devoid of human settlements, contains the maximum 
concentration of this species. In winter the cat is totally absent in sections 
with a snow cover 30 to 35 cm deep, i.e., along mountain slopes 500 to 
600 m above sea level (G.F. Bromlei). Along the coast this cat is most 
frequently met with in burned-over forest sections with 40- to 50-year-old 
Mongolian oak trees and often bushes, hazel shrubs, etc., forming thickets 
impenetrable to man. Manchurian hares and pheasants are abundant in such 
thickets. The cat readily inhabits forest glades also, as well as forest belts 
adjoining cultivated fields, and the shores of marine bays. It does not avoid 
open meadows and untilled lands. 

In all the foregoing situations, mice, field voles, Manchurian hares, and 
pheasants are generally plentiful. Furthermore the snow cover in such places 
is usually not deep and the snow almost throughout winter rapidly becomes 


265 Fig. 162. Herbaceous-shrub growth in the Sukhorechensk range, representing year- 

round habitat of the Amur cat. "Kedrovaya Pad' " preserve near Vladivostok. 
August, 1963. Photograph by A.G. Pankrat'ev. 

covered with a thin crust, facilitating movement of the cat. The predator 
is confined year-round everywhere to cliffs and rock slides, where it sets up 
265 a lair in summer and protects itself from deep snow in winter. Compared 
to the number of Amur cats found in the foregoing habitats, only a small 
population will be found in broad-leaved forests of oak, maple, linden, 
ash, Manchurian nut, philodendron, and others commonly lacking dense 

In nut pine-broad-leaved forests at heights of 300 m above sea level 
the Amur cat is still rarer. Finally, it is absent, as a rule, in the zone of dark 
coniferous taiga, consisting of spruce and fir, that occupies montane slopes 
500 to 600 m above sea level. It usually moves into the taiga only in sum- 
mer and primarily in years when chipmunks and tree squirrels are abundant 
(G.F. Bromlei). Thus, in Suputinsk preserve in the Primor'e, noted for its 
nut pine-broad-leaved forests, this cat appears from time to time only in 
the snowfree period, and confines itself mainly to old clearings where mouse- 
like rodents, chipmunks, and Manchurian hares are abundant (Bromlei and 
Gutnikova, 1955). Consequent to a plentiful snowfall the cat experiences 


a period of food scarcity and descends into the valleys, coming closer than 
usual to human dwellings. Only during periods of heavy snow are cases 
of cat attacks on fowl reported. Its appearance in villages was noted time 
and again, for example, in the very snowy winter of 1935/1936. This cat 
could hardly have survived on chickens, however, since dogs begin to bark 
as soon as the birds squawk (Zubarovskii, 1939; G.F. Bromlei). 

Food. This cat feeds on "mice and various birds, the nests and young 
of which are mercilessly ravaged "; it also attacks hares and young roe deer 
(Przheval'skii, 1870). In summer the predator seeks out nests of marsh and 
water birds and eats the fledglings; lizards and snakes are also consumed. 
In the forest, chipmunks, tree squirrels, and young roe deer fall victim 
(Zubarovskii, 1939). The list of animals preyed upon by the Amur cat in 
Kirin province of China is roughly identical (Baikov, 1915; except for roe 
deer). In years of high populations of mice, voles, and chipmunks, when 
there is no snow, these rodents constitute the primary captures of the Amur 
cat. In winter chipmunks hibernate, and catching mice, especially voles, 
becomes more difficult; hence the cat often resorts to Manchurian hares, 
tree squirrels, pheasants, and hazel grouse. Rarely, it catches small birds. 
In years of low populations of mouse-like rodents, the role of birds in the 
166 diet of this predator increases. Hazel grouse and hare are the main quarry in 
winter. 7Tie cat consumes spawning fish, which it catches along river banks 
and, in winter, in unfrozen ponds and brooks. The cat very rarely attacks the 
young of roe deer, sika, goral, and other ungulates and, when it does, only 
. animals a few days old left unguarded by mother wandering far from them. In 
years of food abundance these cats become extremely obese. Greatly weak- 
ened cats are caught in years of low mouse-like rodent populatioas, especially 
in winter; the stomachs of such cats are either empty or filled with rotten 
wood and tree fungi, evidence of the animal's starvation (G.F. Bromlei). 

In spite of the comparatively small size of this cat, two instances have 
been established of its finishing in a single sitting an entire pheasant, 
including much of the feathers (Bromlei, 1951). Apparently in such cases 
the cat does not actually eat some of the flesh but hides it for later 

Home Range. The size of individual territories in the spring-summer 
period is not known. In spring in Sudzukhin preserve it was roughly estimated 
as 9.0 km^. At the end of winter, when snow is particularly abundant, the 
cat may cover 3.0 to 3.5 km in a single hunt. Once, in March, a cat was 
tracked for a distance of 3.5 km. It made its hunting circuit at night and 
returned to almost the same site from which it left, but entered another 
hollow in a linden tree, remaining there throughout the day. On this hunt 
the cat caught three mice and one hazel grouse (G.F. Bromlei). In winter 
the cat has many shelters in which it rests during the day. 


266 Fig. 163. Montane gulley in " 'Kedrovaya Pad'" preserve. Habitat of Amur cat. 

September, 1963. Photograph by A.G. Pankrat'ev. 

Burrows and shelters. In the Primor'e the den for raising kittens is set 
up in a crevice among rocks and in a hollow. In June, 1936, in the Pos'et 
region, two Amur cat kittens were found in a burrow (Zubarovskii, 1939). 
On the eastern slopes of the Sikhote-Alin this cat lives most often in tree 
hollows (G.F. Bromlei). In northeastern China kittens have been found in 
such hollows (Baikov, 1915). In southern China lairs have been discovered 
in tree hollows and under rocks in thickets (Allen, 1939). 
267 Daily activity and behavior. The Amur cat leads a nocturnal life year- 

round and is seen very rarely in daylight hours because it then rests in its 
den. It is a skillful climber of trees and is usually caught in them when 
trying to escape pursuing dogs. 

Seasonal migrations and transgressions. These aspects have not been 
much studied, but the Amur cat is known to wander. For example, on the 
eastern slopes of the Sikhote-Alin, the snow cover is never very thick in 
the first half of winter. In March, however, so much snow may fall in two 
or three days that it attains a depth of 50 to 100 cm. Given deep, porous 
snow cover, cat tracks will not be seen unless a crust has formed; evidently 
this animal migrates at that time into low-snow areas. 


267 Fig. 164. Forest in '"Kedrovaya Pad' " preserve near Vladivostok. Habitat of Amur 

cat. August, 1969. Photograph by G.N. Simkin. 

In the abundantly snowy winter of 1947/1948, when snow fell in 
November and remained until March, reaching a depth of 85 cm, these cats 
disappeared even from those sections of Sudzukhin preserve where they 
had invariably been found before (G.F. Bromlei). No doubt, in addition 
to migration, some simply died of starvation. 

Reproduction. This aspect is poorly known. In the southern Primor'e 
These animals come into heat in March and live at that time in pairs. Gestation, 
as in other small cats, requires about 60 days. Kittens appear in the second 
half of May (Zubarovskii, 1939). In the Ussuri valley kittens have been 
encountered at the end of April or eariy May (old calendar; Maak, 1861). 
On the eastern slopes of the Sikhote-Alin in recent years animals in heat 
have been sighted throughout the month of March. Kittens appear from the 
first few days of May (G.F. Bromlei). In Sikhote-Alin preserve a pregnant 
female was caught on May 7th^' with eight very large fetuses (V.D. Shemykin 
and Yu.A. Salmin). In Kirin province of China, kittens have been seen in 
May (old calendar) (Baikov, 1915). In the south, in Yunnan province 
(southwestern China), newborn kittens have been found on February 27th 

"Year iiere and below omitted in Russian original — General Editor. 


and April 1st (A. A. Sludskii), and three kittens about two weeks old found 
in southern China on May 29th. Three small kittens were found on Hainan 
Island on March 26th (Allen, 1938). In India kittens are seen in May (Pocock, 

In the Soviet Union litters usually contain two kittens each and more 

rarely one or three (Maak, 1861; Zubarovskii, 1939); evidently larger litters 

268 are possible. This is supported by the capture of the pregnant female 

mentioned above. Litters of this cat discovered in Yunnan province consisted 

of two to four young, and in India, of three or four. 

Growth, development, and molt. The general color tone of the upper 
part of the head, neck, and back in a newborn kitten is light brown with 
a reddish hue. The upper lip is creamy-white. From the upper edge of the 
nostrils a milk-white stripe extends on each side upward through the forehead. 
These stripes are bordered by two blackish-brown lines. On the small muzzle 
and forehead there are consequently two light colored stripes and four 
blackish-brown ones. Along the upper part of the neck and on the back, 
blackish-brown spots are longitudinally disposed on the main background 
color, which fuse into approximately eight parallel stripes. The lower lip, 
underside of the neck, chest, and abdomen are creamy-white, mottled with 
longitudinal blackish-brown spots. The lower part of all four paws is pale 
yellow and devoid of spots. The upper portion of the tail is the same color 

Fig. 165. Female Amur cat, 10 to 11 montlis old, menadng photographer. "Kedrovaya 
Pad' "' preserve near Vladivostok. April, 1%4. Photograph by A.G. Pankrat'ev. 


as the back but slightly lighter and mottled with longish spots. Distally, 
the tail coloration becomes a monochromatic light brown. 

The growth and development of kittens have not been studied. It would 
seem that the young hve with their mother far into the autumn, and possibly 
even until she comes into estnis again, since the tracks of a female with 
kittens were encountered on November 27. The footprints of these kittens 
were almost the same size as those of the mother (G.F. Bromlei). 

Molt in the Amur cat has not been studied. 

Enemies, parasites, diseases, mortality, competitors, and population 
dynamics. No information is available about the enemies of the Amur cat 
except for one report of cat puфortedly torn to bits by an eagle owl [Bubo 

Every cat caught in the spring-summer period, especially June, has been 
infested with mites Ixodes persu lea tus (infestation index 49.3%), Haemo- 
physalis japonica douglasi (48.0%), and H. concinna (2.7%). These mites 
overwinter on cats (up to 40 individuals have been counted in some animals). 
Five species of fleas have been identified: Chaetopsilla appropinguans 
(12.5%), C. micado (18.75%), Paraceras flabellum (6.25%), Ceratophyllus 
tamias (6.25%), and Leptopsylla ostisibirica (56.25%). These fleas were 
detected year-round. P. flabellum is usually specific to badgers, and is 
encountered in them from the Primor'e along the mountains as far as the 
Tien Shan (A.G. Pankrat'ev). In 50% of the cats examined tapeworms were 
present in the anterior part of the intestine, but their species composition, 
was not established. Infectious diseases of Amur cats have likewise not 
been studied. 

In general, competitors of the cat are fox, lynx, and yellow-throated 
marten, since the food of all four is quite often identical. 

Population dynamics have also not been studied. Presumably, popula- 
tioas of this cat would vary significantly in relation to fluctuations in stocks 
of moiLse-like rodents and other animals serving as food. In winters of greater 
than normal snow some cats die from hunger and resultant diseases. In recent 
years, with an increase in areas cleared, expanded forest felling, and 
plantations, the cat population in the southern Primor'e has been increasing 
and the animal is being seen more often in villages (G.F. Bromlei). Present- 
day small-scale hunting has no adverse effect on the cat population. 

Field characteristics. This cat differs from the domestic cat in its much 
larger size, long legs, and comparatively short tail; tail length is equal to 
Z69 one-half of body length with head. The body color is reddish, mottled with 
small rust-brown spots. On the head appear two prominent light-colored 
stripes, running parallel posteriorly along the forehead (see Fig. 165). The 
pace of walking is similar to that of domestic cats, but strides longer. The 
average paw size, measured from tracks, is front paw 37 mm x 38 mm and 


hind paw 36 mm x 30 mm. The cat buries unconsumed prey, for example 
Manchurian hare or fowl, in the forest and invariably returns to it the 
following day (G.F. Bromlei). (A.S.) 

Practical Significance 

The Amur cat is not very important to the fur industry. Throughout its range 
in the Soviet Union, in the best years only 1,000 to 2,000 skins, costing 
1,000 to 1,500 mbles, were tanned. In the Far East area the following number 
of skins were tanned: 1,260 in 1932, 2,177 in 1933, 886 in 1934, and 1,301 
in 1935. Of the 34 regions in which the skins of this cat are tanned, the 
most important are Khankaisk, Grodekovsk, Shkotovsk, and Pos'etsk. In 
each of these regions over 150 skins were tanned in the 1930's, followed 
by the Vladivostoksk, Budennovsk and Voroshilovsk regions, in which 50 
to 150 skins were tanned in a season. Even smaller numbers, i.e., 10 to 50, 
were tanned in the Ol'ginsk, Khorol'sk, Chemigovsk, Ivanovsk, Anuchinsk, 
and Terneisk regions (Zubarovskii, 1939). In the Primor'e territory from 
1935 through 1946 the number of cats caught annually was 104 to 269 (G.F. 
Bromlei). The catch has decreased greatly in recent years, with the total 
output of skins in the Soviet Union placed at only 100 to 300. 

By destroying rodents, chipmunks, and hares, the Amur cat often protects 
crops from pests and hence should be considered a useful animal. It rarely 
attacks chickeas, ducks, and geese, and hence its damage to poultry farming 
is practically nil. It causes more damage to game by destroying valuable 
pheasants, hazel grouse, and fur-bearing animals (squirrels), but the damage 
is not great due to the relative scarcity of the cat. 

According to deer farmers, this cat sneaks into deer parks and destroys 
young sika up to one year of age (Menard, 1930). Referring to information 
obtained from former owners of deer nurseries, G.F. Bromlei (1956) reports 
that this cat attacks sika fawns up to one month of age only when the mother 
is not in the vicinity. 

A forest cat may be caught accidentally during a hunt for other animals. 
Usually it is shot with the aid of dogs, which tree it. It readily enters varioiLS 
traps baited with fresh hazel grouse. 

In the fur industry these pelts are included in the ' 'wildcat" gro p, but' 
identified as "Amur". Formerly, the pelts of this cat were exported to China 
where they were used to make collars, caps, and linings for ladies' jackets. 

The range of the Amur cat in the Soviet Union is small and hence its 
total population quite low. The range is shrinking and the population in many 
parts dropping rapidly; the Amur cat could easily disappear altogether. 
Measures for its protection should be implemented, at least in those areas 
where the animal faces extinction. (A.S.) 



Felis (Felis) chaiis Giildenstaedt, 1776 

1776. Felis chaus. Giildenstaedt. M?v^ Com. Acad. Petrop., vol. 20, p. 483, 

1811. Felis catolynx. Pallas. Zoogr. Rosso-Asiat., vol. 1, p. 23, Terek. 
1898. Felis chaus typica. De Winton. Ann. Mag. Nat. Hist., ser. 7, vol. 

2, p. 291. For F. ch. chaus Giild. 
270 1898. Felis chaus furax. De Winton. Ibid., p. 293, Jericho, Palestine. 
1939. Felis chaus prateri. Pocock. Fauim Brit. India, Mamm., vol. 1, p. 298. 

Jacobabad, Sind, northwestern India. ^^ 
1969. Felis (Felis) chaus oxiana. Heptner. Zoolog. Zhurn., 48, 8, 1259. 

"Tigrovaya Balka" preserve, lower courses of Vakhsk [River], right 

tributary of Amu-Darya. (V.H.) 


Color of trunk and head monochromatic, without distinct spots or stripes; 
dark markings occur only on the tail and legs. Skull long, but palate not 
extended posteriorly in the inteфterygoid vacuity (see Fig. 28); inteфterygoid 
vacuity broad. Second premolar (first in row) usually present; antero-inner 
cusp of camassial tooth well developed and with a high pointed cusp. (V.H.) 


Size small, corresponding roughly to that of Caucasian forest cat [Felis 
silvaticus caucasicus], but somewhat larger in general. Some animals may 
be very large. 

In general appearance jungle cat differs somewhat from forest or steppe 
[wild] cats in that the former stands slightly taller on its legs and has a 
relatively short tail (Fig. 166). Tail length is less than half, usually about 
one-third of body length, and half the length of the hind limb (rarely equal 

^^Also called "chaus'" and sometimes "jungle lynx."* The latter name is artificial, 
simply a translation of the German name, and extremely unsatisfactory since this species 
is not related to lynx. The small tufts on the ears confused earlier investigators. Moreover, 
in old illustrations artists not acquainted with the live animal gave it a lynx-like appearance — 
extremely long legs and lynx-like ears. 

*Also reed cat and reed lynx — General Editor. 

^^sliawiana and maimanah are sometimes included among synonyms for species in 
Russian fauna. These names should not be applied to the jungle cat (see p. 441 footnote 
and synonyms for cats of libyca group). 


to it). These proportions do not, however, reflect a similarity between jungle 
cat and lynx, as often stated. 

Head fairly broad in region of zygomatic arch and hence appears more 
rounded (spherical) than that of Amur cat. Ears quite long, relatively broad 
at the base, pointed toward the end, and set quite high. Small tuft of long 
hairs occurs on ear tips in winter. These hairs form an indistinct tassel ranging 
from 7 to 10 to 20 mm in length. Ears invariably project out of coat. Hair 
on cheeks not long (side whiskers not developed). Eyes fairly large and pupil 
vertical. Head or "face" of jungle cat altogether typical of small cats, and, 
like that of the forest [wild] cat, has nothing whatsoever in common with 
the lynx. 

Pelage throughout the body uniform in length, but in summer, hair on 
back long and forms a narrow and sparse crest of bristly guard hair all along 
the spine. The hair in this crest may be up to 50 mm long. In some animals 
this crest is weakly developed, however. In winter coat a crest per se is 
not seen, but the hair along the middle of the back is perceptibly longer. 
It is quite possible that the summer crest is formed of unmolted winter hair. 

The guard hair of the winter coat is relatively long and resilient and 
hence, in spite of the soft underfur, the coat of a jungle cat, compared to 
that of other small cats, is sparser and rougher. Neither in winter is it long 
and rich, but sparser, short and rough, as in summer. In winter about 4,000 
hairs occur per cm^ on the back and 1,700 on the abdomen. Extreme growth 
of top hair on the abdomen is a characteristic feature. The ratio of underfur 
to top hair on the back is approximately 12 : 1 and only 4 : 1 or 5 : 1 on 
the abdomen. The top hair is very long; that of category I is 1 .8 times longer 
than the fur. The average length of guard and top hairs of categories I to 
IV on the back is 61, 56, 47, 43, and 39 mm, and their thickness 89, 74, 
272 52, 40, and 30 microns^^; on the abdomen the corresponding values are 50, 
37, 30, and 26 mm (top hair of category IV absent), and 48, 30, 28, and 
27 microns. The underfur is 30 mm long and 20 microns thick on the back 
and 20 mm and 21 microns on the abdomen (B.F. Tserevitinov). 

On the whole the body color of a jungle cat is monochromatic and 
devoid of spots. Dark markings and spots occur only on the legs and tail and 
sometimes on the abdomen. The general color of the winter coat (average 
type of coloration of Caucasian animals) is grayish-ocherous with an 
indistinct, very slight ripple consisting of light and dark top hairs. On average, 
more white hairs occur in this "ripple" and hence the coat has a grayish 
cast. Behind the scapular region a band extends along the back, which is 
initially narrow and indistinct but later becomes quite broad and turns a 

'"^Measiirements along the middle of the back, i/e., in the region of the "crest", are 


deeper reddish-ocher color. Dark-colored hair is finely sprinkled in this 
band near the scapular region, but its density increases toward the rump. 
On the sacrum, and sometimes at other places, the density of dark-colored 
hair may actually form narrow fringes along the ocherous band. 

Flanks lighter in color than the back and admixture of top hair (small 
ripple marks) reduced; down the flanks, color merges rather abruptly into 
that of abdomen. Chest and stomach a vivid light ocherous shade with 
sections of white on chest and axilla; posterior third of abdomen and groin 
regions white. Chin, region of lower jaw, and throat also white. Basal part 
of neck and anterior part of chest a vivid light ocherous color. This field 
expands in the form of a cape into the region between the forelimbs. 

Outer side of hind legs identical in color to the flanks; outer side of 
forelimbs dull ocher; admixture of dark top hair in latter less than on hind 
limbs, or absent, and hence forelimbs appear brighter than hind ones. Two 
dark brown transverse stripes run into upper portion of inner side of forelimbs 
on an extremely light ocherous -white background, or a few spots to five or 
six short stripes of the same color may occur. Top of neck the same color 
as anterior part of back; sides of neck slightly lighter and approach color 
of the flanks. Uppermost part of tail the same overall color as the back 
or sUghtly grayer — ocherous stripe reaching base of tail does not extend onto 

Fig. 167. Young jungle cat. Trans-Caucasus. Photograph by A.P. Zhandarmov. 


it. Proximal portion of tail exhibits a strong admixture of black top hair; 
tip black with two or three black rings above it; first and second rings 
sometimes connected by a longitudinal band. 

On the head, between the ears and the occiput, color the same as on 
>73 the back but more saturated and brownish. Upper lip and region of vibrissae 
bright white, or slightly ocherous, and stand out (characteristic features of 
the species) (Fig. 168). A dark brown spot occurs in front of the eye toward 
the nose, and a rather vivid ocherous field under the ear, behind it, and on 
the cheeks. Rest of the head same color as the flanks. Back of ears a vivid 
ocherous-red, but slightly brownish at the base; tip of ears brown or black, 
and tuft black. Inner side of ears covered with white hairs, the longest of 
which are pure white. Vibrissae white. 

Individual variability of winter coat relatively minor; coat either more 
grayish or more ocherous in general shade, and shape and color of band 
on back varies. Latter may be short or long, very broad and indistinct, or 

Fig. 168. Jungle cat. Felis {Felis) chaiis Giild. Bright coloration of upper lip a 
characteristic feature. Photograph by M.l. Obukliov. 


relatively narrow, зЬаф, and rather vivid. Intensity of color of the underside 
of body also varies, but bright spot on neck and chest distinct. Brightness 
of color and outlines of spots vary slightly. Large number of spots (bands 
on forelegs invariably two), and poorly developed, indistinct ocherous spots 
may be visible at the boundary between color fields of flanks and abdomen, 
as well as on the abdomen. 

Color of summer coat far more variable than that of winter coat. On 
the whole it is far brighter and only rarely as dark and dense as the winter 
coat. General color tone of back varies from ocherous-gray to almost pure 
gray, with a slight dark- and light-colored ripple — gray tone may be quite 
dark or very light. Stripe on back, as in winter coat, varies in length and 
width and ranges from ocherous-brown to bright ocherous-rust. Color of 
underside of body less vivid and far lighter — white fields larger and ocherous 
tone paler. Sometimes, however, entire underside of body, except for chin, 
a uniform bright ocherous color, even exhibiting an orange hue. Spottedness 
somewhat more intense in summer than in winter; spots on legs and at border 
of color of flanks and underside more vivid, while spots on abdomen better 
developed and sometimes fuse into transverse stripes. 

Very light-colored and pale animals^^ obviously represent an extreme 
and very rare type of color variation in the summer coat. Top of neck light 
and grayish; only a relatively narrow gray band (almost equal in width to 
ocherous belt) extends down the back; sides, however, light ocher. Color 
may also be ocherous-gray with a rusty band. A narrow black stripe occurs 
on the tail which, as such, is white. Lower portion of tail very light-colored. 
Chin, throat, and chest between legs white, and area between chest and throat 
light ocher. Posterior part of chest and anterior part of abdomen pale ocher, 
274 and posterior part of abdomen and groin white. Spottedness noticeably rich. 
They are well expressed along legs and abdomen, or legs, flanks, whole 
of abdomen, and whole of chest covered with rust-brown (hind legs and 
flanks), brown (forelegs), and bright rusty (flanks and underside) indistinctly 
defined spots. Coloration in this type of spotted pattern is quite dramatic. 

General color of first coat in young animals (June, Kushka) dirty gray. 
A broad, poorly delineated dark brown-gray field extends down the back. 
Along the sides of this field, on the back, flanks, and abdomen, indistinct 
gray spots tend to form vertical rows. The field between them on the back 
and flanks is a slightly lighter gray. Underside of body light, pale yellowish- 
white, with yellow hue deepening in anterior part of abdomen and on the 
chest. Forelegs above the ulna rust-colored but dull; small blackish-brown 
spots occur below the ulna on both the outer and inner sides. Hind legs bear 

-''No. S 14194; Alebandriisk station, lower Terek; July 5, 1926; Yu.G. Heptner. No. 
S 52701; Gasan-kuli at mouth of Atrek, Turkmenia, 1942; A.V. Samorodoy, KZMMU. See 
later section, "Systematic Position" 


brown spots and both upper and lower parts of feet rusty. Dorsal side of 
tail dark brown and underside light ocherous. Top of head same color, as 
the back and sides slightly paler. Field under the eye and to the rear of it, 
under the ear, ocherous-gray; upper and lower lips, chin, and throat white 
(see Fig. 167). Back of ear rust-colored, tip black, and ear tuft black and 
about 10 mm long. Juveniles assume an adult coat in their first winter. 

This species displays distinct geographic variation in coloration and 
is a good example of Gloger's rule (Weigel, 1961). In addition to a general 
intensity of color in some subspecies, range and type of individual variation 
also seem to differ — gray and reddish-brown phases, together with 
transitional colors evident in the torso (F. c. ajfinis), darkening of color of 
the legs, and so on. Within the Soviet Union color variation is insignificant 
and mainly reflected in general hue and sometimes individual patterning. 

The skull of the jungle cat (Fig. 169) occupies an intermediate position 
between that of the peripheral form (in the Soviet fauna) of the genus — the 
Amur cat — and more specialized forms of the subgenus Felis s. str. — group 
silvestris, species silvestris. This similarity is exhibited in the slightly longish 
form of the skull in general and the brain case in particular, the comparatively 
narrow zygomatic arches, the form of the nasal processes of the premaxilla, 
and in other details. At the same time, the brain case in jungle cat is more 
expanded, roomier, and less extended posteriorly, the frontal area large, 
bulging, and with a small depression in the center, and the entire skull per 
se more convex (upper line of profile even above the arch is acute). 
Furthermore the orbits are relatively and absolutely larger; though set only 
slightly more forward, they extend slightly more posteriorly and take on 
an oval shape. The palate does not extend posteriorly to the same depth into 
the interpterygoid vacuity as it does in the Amur cat; in the jungle cat the 
posterior boundary lies at the level of the posterior margin of the molars 
(see Fig. 28). The interpterygoid space is broad, somewhat broader in front 
than behind, and the width of the choanal opening much greater than its 
height. The anterior width of the inteфterygoid vacuity exceeds the distance 
between the inner margins of the auditory bullae. 

The teeth, especially the carnassial and third premolar, are very strong 
and massive. The anterior inner cusp of the carnassial tooth is well developed, 
with a fairly large pointed peak. The second premolar is usually present. 

Most of these features are developed to such an extent that in differentiat- 
ing the jungle cat from the Amur cat, the former comes somewhat closer 
to forest and steppe [wild] cats, and even to the caracal (see "Systematic 

Age-related changes in the skull are evident in the relatively larger 
volume of the cranium, lesser expansion of the skull per se and its more 
spherical shape, lesser development of the orbital processes, and absence 


or poor development of the sagittal and lambdoidal crests. The proportions 
and structure of the skull of an adult animal are not seen in a juvenile prior 
to the middle of its second year (which means its second winter). The skull 

275 of females are usually lighter and somewhat smaller than that of males. 
Geographic variation in the skull is almost nil and seen only in insignificant 
differences in overall size. 

In body size, jungle cats in various parts of the Russian range do not 
vary much, and the data given for the region in the upper reaches of Amu- 
Darya (Termez and above. Table 7) are also applicable to the Caucasian 

Significant variations in weight (twofold or more in males and almost 
threefold in females) are attributable not only to significant individual and 
age-related variability, but also to seasonal changes and availability of food 
in different years. In the Amu-Darya delta the weight of males (13) in 
March-April averaged 7.9 kg and in October to December (4) 8.2 kg; the 

276 corresponding values for females (4) were 5.5 and (3) 6.5 kg (Reimov and 
Nuratdinov, 1970). Under favorable conditions these cats can become ex- 
tremely obese. 

Absolute mean weight of heart in males (21) 39.2 + 3.9 with a coefficient 
of variation of 34.2 and in females 35.8 ± 4.7 and 29.3 respectively. Relative 
weight averages 4.7 ±0.25%o with a coefficient of variation of 17.0 and 
6.1 ± 1.0 and 36.7 respectively. Absolute average length of intestine in males 
(21) 183.9 ±5.9 (coefficient of variation 10.98) and relative length (for 
average body length of 76.5) 1 : 2.40; absolute length in females (15) 
176.5 ± 12.0 (coefficient of variation 10.82) and relative length (for average 
body length 68.8 ± 2.2) 1 : 2.56. The absolute and relative weights of the 
heart, depending on degree of obesity, also reveal quite significant individual 
variations, often in different seasons of the year — males averaged from 
39.7 to 34.6 g and 5.02 and 4.22%o, and females 40.4 to 29.2 g and 7.34 
and 4.46%o (according to Reimov and Nuratdinov, 1970 with modificatioas; 
data for Amu-Darya delta). 

Skull measurements, based on data for the whole of the Russian range 
of the species, except the Amu-Darya delta, are given in Table 8. Data for 
different sections of the range are presented under "Geograpliic Variation". 

As in the case of body size and weight, the skull measurements of males 
significantly exceed those of females by an average of 15 to 16% with respect 
to the first category of characteristics, and roughly by 10% with respect 
to the second category. (V.H.) 

277 Systematic Position 

As briefly mentioned above, the jungle cat in skull structure (see Fig. 169) 


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115 Fig. 169. Skull of jungle cat, Felis (Felis) chaus Giild. No. S 5376, male, Zoological 

Museum, Moscow University. Lenkoran, Vel', February 10, 1925. Sketch by 

N.N. Kondakov. 


occupies an intermediate position between F. euptilura andF. silvestiis — 
libyca, i.e., European forest and steppe cats. In this species (superspecies) 
F. silvestris, the jungle cat is closer to the subspecies group sllvestris than 
to libyca. Present-day taxonomists usually place it in subgenus (of genus) 
Fells s. str. (Pocock, 1939; Haltenorth, 1953; and others). In spite of 
prevailing views, in this subgenus the jungle cat is a less specialized form 
(slightly extended skull, cranium, position of orbits, and so on). This fact 
also determines its position outlined above, although the structure of the 
palatal, choanal, and inteфterygoid regions differentiates the jungle cat from 
the Amur cat more than from European forest and steppe cats (species - 
superspecies sllvestris). 

The indisputable taxonomic proximity of the jungle cat to the latter 
does not permit its segregation into a special subgenus {Chaus) as done until 
recently and sometimes even now (Petzsch, 1968). In view of the reasons 
stated above, there is no justification for bringing the jungle cat closer to 
lynx, ahhough, as will be shown below (in section on caracal), in certain 
features of skull and level of its specialization from the Amur cat — manul 
lineage, the lynx (subgenus Lynx) can be separated from subgenus Fells at 
the moфhological level of the jungle cat. In general body build, which reflects 
basic features of the lynx group, the jungle cat nevertheless represents a 
true cat of the steppe or Amur cat type (traditional opinions about this animal 
in this respect are biased). 

The color of the jungle cat is distinctive and differs markedly from that 
of other cats of the narrow subgenus as inteфreted here. However, the spotted 
color of the young and individual variations in the spotted pattern of adults 
point to the close relationship between the jungle cat and the species of true 
cats. An intermediate color type, between the Amur and European forest 
cat, is thus not seen, ahhough the similarity of the monochromatic color 
type of the Amur cat [see above] to the color of the jungle cat is quite evident. 
It is significant that in captivity (zoological parks), jungle cats produce fertile 
offspring with domestic cats (Gray, 1954); a more extensive analysis of 
hybrids has not been made. 

There is less definite information about the mating of wild Indian jungle 
cats with domestic cats (Allen, 1938). Nothing is known about this in the 
Soviet Union. 

The above-mentioned differences between jungle and Amur cats in fact 
represent a unique set of distinguishing features between Amur and forest 
cats. Hence, with the information available at present, there is no justifiable 
basis for separating F. euptilura from cats of the subgenus Fells s. str. It 
is possible that these differences also may be found in other species usually 
placed close to F. euptilura {bengalensls, vlverrlna, rubiginosa, and 
planlceps — Prionailu7-us group). 


The color and some features of the build of the jungle cat are distinctive 
and have no transitional types. The position of the jungle cat is also important 
in that it bears significant features of similarity with caracal in craniological 
features. Being a wholly typical member in the direct line of specialization 
of Soviet cats from Amur to manul, it represents that level in this chain 
of forms from which, so to speak, the lynx (subgenus Lynx) branched off 
(see characteristics of the genus and "Systematic Position" of caracal). 

Geographic Distribution 

Jungle cats are found in forest regions and areas close to water in the Near 
East, Middle, and Southern Asia, and northeastem Africa. 

Geographic Range in the Soviet Union 

The range in the Soviet Union (Fig. 170) represents the northern periphery 
278 of the range of the species. It covers the southernmost regions of the country 
and is divided into two parts — Caucasian and Middle Asian — which unite 
in the south outside the Soviet Union. Because of its preference for river 
valleys and other areas with abundant water, often with forests, the distri- 
bution of the jungle cat is extremely uneven and boundaries of its range, 
especially in the Soviet Union, are very complex. Extensive areas in which 
the animal is absent have been delimited. 

In the northwest jungle cats live in the Volga delta at places 65 km 
north of Astrakhan (Dosang) and among the Pri-Caspian reeds for a bit 
east of the delta (Dengizsk region, Guryev district). From the Volga delta 
the range extends southward to the mouth of the Kuma, and along its lower 
reaches, in the form of a narrow strip corresponding to the width of the 
reed beds along the Caspian coast. 

From the mouth of the Kuma the cat is distributed along coastal reeds 
southward to Chemyi market. It inhabits floodplains and stream channels 
in the area between the Terek branch running north to Chemyi market and 
branches [of the delta] running east into Agrakhansk Bay. Along the Terek 
it extends into the forests and thickets of the upper valley as far as Parabochev 
forest and Shelkozavod (Shelkovaya) station, and even to Mozdok and along 
the Sunzha to Groznyi. In the expanse between the Terek, in the foothills 
[of the Caucasus] and on the seacoast southward to Makhachkala (Dagestan 
plains) the cat is quite uniformly and extensively distributed, some distance 
from the sea, along the floodplains of the Terek, Sulak and its tributaries, 
and along lakes and shrubby thickets. It reaches as far as a line between 
Shelkovaya and Khasavyurl, or perhaps even farther westward. 

• 369 

279 To the south of Makhachkala the jungle cat is found along the Caspian 

coast to the Iranian border. Where reed thickets are absent this cat lives in 
shrubby thickets in the coastal belt or in foothills, rising up to heights of 
300 to 400 m, and not more than 800 m, above sea level, but up to 1,000 m 
in the Trans-Caucasus. 

This cat is extensively distributed in the lowlands of the eastern Trans- 
Caucasus, along the Kur' and Araks valleys and Kur' tributaries, and among 
lake regions in the steppes. The range there is bounded by the foothills of 
the Great and Little Caucasus. In the Kur' valley it extends almost to the 

278 Fig. 170. Reconstructed range of jungle cat, Felis (Felis) chaus Giild. in the Soviet 

Union (scale in km). Question marks pertain to the lower Chu and the Ferghana 
valley, information concerning which is dubious. V.G. Heptner. 


Suram range (Gori) along the lower zone of the mountain and in the foothills 
of the Glavnyi [Caucasus] range and the Alazan basin, and runs in the west 
roughly to the meridian of Tbilisi. Along the river valleys of the Kur' basin 
extending down from the Little Caucasus (Terter), it extends at places quite 
deep into the mountains. The cat lives in the Araks valley from its lower 
reaches up to Megra ravine (a spur of the Zangezur range on the Araks, 
slightly east of Ordubad) and, after several breaks (southeast of Yerevan 
at the boundary of Nakhichevan Autonomous Soviet Socialist Republic), 
from Arazdayan to the Oktemberyansk region (west of Yerevan). It even 
penetrates there deep into the mountains along some tributaries (Akera and 

References to the occurrence of the jungle cat in the Kuban (M. 
Bogdanov, 1873; Ognev, 1935; Formozov, 1946) are erroneous; only the 
forest [wild] cat (F. silvestris^^) occurs there. Information about its 
occurrence at the mouth of the Ural [River] has not been confirmed and 
is evidently erroneous. 

In Middle Asia the jungle cat lives in the extreme southeastern comer 
of Turkmenia along the Atrek from its mouth (Gasan-kuli) and its tributaries, 
the Sumbar and Chandyr, and along the Tersakan (Khodzha-Kala). The 
animal does not penetrate deep into the mountains (along the Sumbar hardly 
beyond Kara-Kala). Throughout the rest of the Kopet-Dag this cat is 
evidently absent and also not seen in the adjacent plains. It does not go 
into the [desert] sands and references to its occurrence west of Uzboi 
(Yashkha and Toniatan; G.P. Dement 'ev, 1955) are erroneous. Farther east 
the cat is found all along the Tedzhen, probably entering from Iran, and 
the entire Murgab. It is absent or rare along the Kushkha. 

In the Amu-Darya deha this cat is distributed to the [Aral] sea, and 
up the river to the mouth of Kyzyl Su and slightly beyond (to Chubek, 
south of Kulyab), i.e., all along the river wherever reeds and shrubby 
thickets grow. Along the left bank of the river, beyond the delta, the cat 
is encountered only in its valley (desert closely approaches it). Only along 
the Kelifsk Uzboi and at the head of the Karakum canal does it deviate 
slightly from the Amu-Darya valley (probably the canal later leads the cat 
deep inside the desert). 

The jungle cat is extensively distributed along the right tributaries of the 
Amu-Darya; along the Kyzyl Su and its tributary the Yakhsu the boundary 
proceeds slightly above the Kulyab, runs along the Vakhsh a little above 
Kurgansk-Tyub (where the river flows out of the mountains), and along the 
upper Kafirnigan. It reaches the Gissar valley, in the east slightly beyond 

-^A specimen from Grivensk station (Zoological Museum, Mascow University), labeled 
jungle cat, belongs to this species. 


Dushanbe (to Ordzhonikidzeabad), and along the Surkhan-Darya valley and 
the lower Tupalang. In the mountains between the above-named rivers it 
does not occur. Probably it lives along the Shirabad. It is distributed along 
the Zeravshan, where it is met with especially around Samarkand, and the 
boundary mns upstream along the river at least to Pendzhekent; it evidently 
lives on the Kashka-Darya. The animal occurs throughout the Syr-Darya, 
from the mouth of the Angren at the [Aral] sea, throughout the lower Chirchik 
and Angren, and the plains of the Tashkent region. Data are not available 
for the Ferghana valley. The cat apparently occurs along the lower reaches 
of Chu but these data require confirmation. 

References to the occurrence of the jungle cat in Ustyurt (Brandt, 1852; 
Bazhanov, 1951), not only in the central region, but also between the northern 
Chink and the Bol'shoi Barsuk sands, are erroneous. The cat lives around 
Lake Sudoch' in the extreme northwestern part of the Amu-Darya delta, 
and short transgressions northward from there along the west shore of the 
Aral Sea should not be totally ruled out. 

The range of the jungle cat, in spite of the animal not being subjected 
to special intensive pursuit, and being known to survive well under certain 
280 conditions in cultivated areas (flood- and sprinkler-irrigated lands) undergoes 
considerable changes in certain places. This is explained by its characteri- 
zation as a stenotopic species. In recent years, as a result of the drying up 
of the Atrek and the disappearance of reeds along the river and associated 
water sources (Lake Delili), the entire reed bed fauna has vanished from 
this area, and with it the jungle cat. 

In southern Tadzhikistan the cat has disappeared at several places as 
a result of clearing of the tugais [bottomland forest and shnibland]. This 
has brought about changes in the distribution of the species along the Murgab. 
Contrarywise, as a result of irrigation and the cuhivation of some regions 
in Tadzhikistan which until recently were deserts the jungle cat has reap- 
peared for example in the Vakhsh valley. 

There is reason to believe that the jungle cat, a normal inhabitant 
of the whole of the Syr-Darya valley, disappeared in the last decade from 
its lower course, or at least became rare, even though changes in the 
environment did not occur which would preclude its existence in this 

"Range based on data of M. Bogdanov, 1873; Zanidnyi, 1890 and 1915; Dinnik, 
1910-1914; Satunin, 1915; N. Smimov, 1922; S. Naumov, 1927; G. Nikol'skii, 1930; Ognev, 
1935; Flerov, 1935; Shnitnikov, 1936; Gureev, 1937; Dobrokhotov, 1939; Heptner and 
Formozov, 1941; Vereshchagin, 1947; Kuznetsov, 1948 and 1948a; Chemyshev, 1948 and 
1958; Bazhanov, 1951; Dal", 1952 and 1954; Sludskii, 1953; Heptner, 1956; Nur-Gel"dyev, 
1960; Shukurov. 1960; Ishunin, 1961; Alekperov, 1966; O.P. Bogdanov, 1964; and others; 
and original data of A.A. Sludskii and V.G. Heptner (Turkmenia). 


Geographic Range outside the Soviet Union 

The range outside the Soviet Union (Fig. 171) covers lower [elevations of] 
Afghanistan, Iran, Asia Minor (west and south), Iraq, parts of Syria, Palestine, 
Egypt (Nile delta and the lower course to Kep at 28° N. lat. and along 
281 the coast for 250 km west of Alexandria), and also Baluchistan, Kashmir, the 
whole of India and Sri Lanka, Nepal, Burma, Indochina (but not Malacca), 
and Yunnan.^"^ 

Beyond the limits of its present range the jungle cat is known from the 
Holocene deposits in Germany and Switzerland (?), among relicts of the 
Azilian and Tardenoisian periods in the Crimea (Shan'koba) (V.I. and V.I. 
Gromov, 1937, after Pidoplichko, 1951), and in Slavic settlements of the 
eighth to the thirteenth centuries, in Pohava territory (I. Gromov, 1948). 
That these remains belong to the jungle cat has justifiably been doubted 
(Vereshchagin, 1959). The identification of a species is hardly possible from 
such remains as were recovered. No other data are available about the 
occurrence of the jungle cat in the southern European part of the USSR. The 
jungle cat, at least in the Caucasus, is evidently a recent immigrant from 
southern Asia (V.H.). 

Geographic Variation 

Geographic variation in the jungle cat, in spite of its relatively small range, 
is quite considerable. Usually eight subspecies are recognized (Ellerman and 
Morrison-Scott, 1951 and 1966), QxcXvsSmgmaimanah (see below; Pocock, 
1951); or with the inclusion of the irregular, recently described form 
valbalala (Sri Lanka — 9 (Weigel, 1%1). Differences mainly pertain to color 
(its overall intensity, color of extremities, degree of variability, and so on), 
which conforms to Gloger's rule (Weigel, 1961); also, extremely massive 
teeth have been described in one form (fiirax) (Pocock, 1939) but this feature 
is not fully convincing as judged from the material available with me. There 
obviously exist some differences in the overall size of some of the subspecies. 
For Soviet forms Bergman's rule is also applicable. In spite of all this the 

■^-'By placing F. shawiana among the forms of F. chaiis, Pocock (1939) included 
Chinese Turkestan ( Yaikand) in the range of the jungle cat. In this respect he was followed 
by some western zoologists in particular Ellerman and Morrison-Scott (1951). This 
misplacement, as mentioned before (see synonyms of jungle and steppe [wild] cats), is 
evidently based on a misundei-standing of the data of collections, since jungle cats do not 
occur farther east than the Syr-Darya and perhaps the Chu. There is no positive information 
about the occurrence of this species in Yarkand; its occurrence there is highly improbable. 

Jungle cats have reportedly been sighted in northwestern Africa (Heim de Balzac, 1936); 
this information is not confirmed. 


280 Fig. 171. Species range of jungle cat, Felis (Felts) chaiis Giild. (scale in km). 

Question mark refers to lower Chu. V.G. Heptner. 

distinguishing features of some subspecies have not been clearly defined 
and hence the actual number of geographic forms will probably prove fewer 
in future studies. 

Two subspecies are known in the Soviet Union. 

1. Caucasian jungle cat, F. (F.) с chaus Giildenstaedt, 1776 (syn. 
catolywc, typica). 

Color relatively dark and traces of a spotted pattern relatively distinct 
("Description"given above pertains to this form). 

Large-sized form. Adequate data oh body size and weight are not 
available but, judging from skull size (Table 9), they are not inferior to 
the form oxiana (Table 8). The weight of three adult males was 6,200 (June), 
7,400, and 5,700 g; two females weighed 5,560 and 6,800 g (Azerbaidzhan; 



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A. A. Sludskii); an old female (Astrakhan' preserve) weighed 13 kg 
(Dobrokhotov, 1939). 

This subspecies is found in the Volga delta, the Caucasus, and Kopet- 

No extension of the range eastward into Turkmenia is known. Animals 
from Kopet-Dag (KZMMU) in winter coat are somewhat lighter than 
Caucasian animals and present a somewhat transitional condition between 
this and the next subspecies; they should nonetheless still be considered 
the nominal form. 

Outside the Soviet Union this subspecies lives in Iran, Turkey adjacent 
to the Trans-Caucasus, Iraq (Mesopotamia), Syria and parts of Turkey adjoin- 
ing it, and Palestine (?). 

The nominal form is distinguished from almost all the other subspecies 

282 by its much larger size. In all the forms except fu?-ax and nilotica, the 
greatest length of skull from the published, though scanty, data (Pocock, 
1939 and 1951) does not exceed 117, 120, and 127 mm respectively, 
and the body length of one does not exceed 80 cm. However, for the 
form furax (Iraq, Syria, and Palestine) the greatest length of skull attains 
131 and even 138 mm. Evidently it is identical with the nominal form; 
moreover most reports are from places quite close to the area where 
the nominal form is distributed (Baghdad; Pocock, 1961), and a large 
part of its range lies in the same natural region of the Near East as 
does the Caucasian form. The form from the lower Nile (nilotica — 131 mm) 
is also evidently close to the nominal form in size but will not be discussed 

2. Turkestan jungle cat, F. (F.) c. oxiana Heptner, 1969. 

Color of trunk lighter than that of nominal form, with a lesser 
growth of black top hair, a more vivid band on the back, and more white 
hair on the underside. Underside of flanks very bright and ocherous, 
spots on legs fainter, and traces of spots in other parts of the body less 

Large-sized animals, evidently not smaller than the nominal form (see 
Tables 9, 10, and 11). 

283 The average body length of adult females from the Amu-Darya delta 
(15) is 668 ± 22 mm and average weight (15) 6.08 ± 0.13 kg (Reimov and 
Nuratdinov, 1970). 

The relationship of this form to that described from India is not 
clear. It is possible that it may be close to the iormprateri, which occupies 
Sind and the region southeast of it. The two are hardly identical, how- 
ever. The type locality of this form (Dzhakobabad) is a straight-line distance 
of about 1,000 km from Termez on the Amu-Darya and beyond Hindu Kush. 
Furthermore the nominal subspecies is found in Iran and Baluchistan 




























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OS о On On 




















1— ( 
































^ i! ?> 

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283 Table 11. Average body size (mm) and weight (Tvg), and skull measurements (mm) 
of male jungle cats, F. (F.) c. oxiana from Amu-Darya delta (from Reimov and 
Nuratdinov, 1970, with modifications) 

Indexes n Males 

Greatest skull length 17 123.8 ± 2.06 

Condylobasal length 17 112.9 ± 1.84 

Zygomatic width 17 81.0 ± 1.46 

Interorbital width 17 20.9 ± 0.46 

Body length 21 755.0 ± 32.00 

Weight 21 8.9 ± 0.56 

Note: The average skull measurements of females published by these authors are not 
given since they are clearly exaggerated and not representative due to the small size of the 
sample (five animals) (V.H.). 

(Pocock, 1951). Moreover, the Sind form is notably smaller than the 

The following forms are usually recognized from some parts of the range 
outside the Soviet Union: 1) F. (F.) c.jiirax de Winton, 1898 — Mesopotamia 
and Syria (see above under the nominal form); 2) F. (F.) c. nilotica de 
Winton, 1898 — Egypt; 3) F. (F.) c. affinis Gray, 1830 — Himalayas from 
Kashmir to Sikkim; 4) F. (F.) с kiitas Pearson, 1832 — northern India from 
Bengal to Kutch; 5) F. (F.) c. prateri Pocock, 1939 — Sind and the area 
to the southeast; 6) F. (F.) с kelaarti Pocock, 1939 — Sri Lanka and southern 
India south of Krishna River; 7) F. (F.) с fulvidina Thomas, 
1928— Indochina and Burma. (V.H.) 


Population. The jungle cat is rare in the extreme north of its range, in the 
Volga delta. It is rare and sporadic on the northwest coast of the Caspian 
Sea. On the lower Terek and southward, to the boundary with Iran, it is 

^The name maimanah Zukowsky, 1915, assigned on the basis of a visual examination 
(!) of the skin from Maimana in northern Afghanistan described by Scully in 1887, could 
probably be given to this form on the basis of marked similarity. Ellerman and Momson- 
Scott (1951 and 1966) assign this name to the jungle cat with doubt (?). However, it refers 
to the steppe cat (see p. 442). 


common, and at places abundant (lowland regions of Dagestan and Azer- 

284 baidzhan, and in parts of Georgia and Armenia). In Azerbaidzhan, until quite 
recently, some 3,500 of these cats were caught annually. The animal is now 
common but was formerly abundant at places in the southern half of 
Turkmenia (along the Atrek, Sumbar, Tedzhen, and Murgab). It is abundant 
on the Amu-Darya and its delta and on the southern coast of the Aral Sea. 
It is common and at places abundant in the tugais of the Pyandzh, Vakhsh, 
Kyzyl Su, Kafirnigan, and other tributaries of the upper Amu-Darya. It is 
particularly abundant on the Amu-Darya along the Afghan border. In 
Tadzhikistan up to 300 cats are caught annually. 

In Uzbekistan jungle cats are extremely common in the valley of the 
Amu-Darya and its tributaries (Surkhan-Darya, Kashka-Darya, and others) 
and in the Zeravshan valley. In recent years it has been rare in the Syr- 
Darya, Chirchik, and Angrensk valleys. In Syr- Darya and Chu in adjacent 
Kazakhstan this species is no longer met with, although earlier authors 
considered it common there. 

Habitat?^ The jungle cat is a typical inhabitant of reed thickets, im- 
penetrable thorny bushes, dense, wet lowland forests around lakes, river 
floodplains, and along sea coasts. In open steppes or deserts it is encountered 
rarely and never ascends high into the mountains. The above-mentioned 
regions abound in swamp and water birds, francolins [Francolinus], water 
or complex-toothed rats [Nesokia iiidica], mouse-like rodents, several species 
of reptiles and amphibians, and also fish. The cat is assured of abundant 
prey in these hunting grounds. In the Volga delta it lives in reed and cattail 
thickets, which alternate with ilmens [Volga delta lakes], and are dissected 
by abundant small rivulets — eriki Willow shrubs grow on the banks of 
these shallow channels. While hunting, the cat sometimes reaches the coast. 
It lives in reed and cattail thickets with a large number of breaks in the 
lower Kuma, Terek, and other rivers of the Caucasus and Trans-Caucasus 
and also in the large system of lakes (Sary Su and others). In Armenia the 
animal lives, for example, in the Araks valley in and adjacent to reed shrubs 
at a height of 800 to 970 m above sea level (Dal', 1954). 

On many of these water sources, reeds and cattails grow in the water 
in the form of a strip 1.0 to 3.0 km wide. These bushes surround the pools 
like a dense wall, isolating them from the adjoining steppes. There are often 

285 bogs and floating islands in these lakes. The cat lives in such thickets 
year-round and ventures into adjoining sections only occasionally and 
briefly, for example the steppe. Hither it sometimes emerges in winter to 
hunt rodents and steppe game. This cat when hunting enters the water and 

•^Photographs given before of habitats of Turanian tiger, except for Semirech'e (Hi 
River) also pertain to this species. This cat and tiger are sympatric almost everywhere. 


284 Fig. 172. Habitat of jungle cat in Astrakhan preserve. Volga delta. June, 1950. 

Photograph by A.A. Sludskii. 

is an excellent swimmer. In the Trans-Caucasus it is often found in breaks 
in the reeds and bogs 1.0 to 1.5 km from the bank. It swims through large 
pools not only in summer but often even in winter. It is so accustomed to 
water that it often rests in breaks among reeds far away from the bank and 
sometimes even sets up its den there. When chased by man or dog, it usually 
jumps into the water and escapes by swimming. 

In addition to reed thickets jungle cats often live in almost impenetrable 
growths of blackberry and clematis growing near water, and in tugai forests 
of turanga [Euphrates poplar] and blackberry. In tugais it is quite common, 
for example on the Kur'. In the Lenkoran lowland and also in other places 
it is not infrequent in the shrub zone of the foothills and in reeds by the 
sea. On the coast it lives in winter wholly on emaciated ducks and coots 
and also on injured animals. Tracks of this cat are encountered everywhere 
along the outer edge of the thickets and on open sandbars. Here the remains 
of its prey are often encountered, i.e., birds torn to bits. (Yu.A. Isakov). 

In the Lenkoran lowland, however, cats are abundant in gardens, where 
they live in hedgerows consisting of white willow densely intertwined with 
climbing, thorny vegetation. At places these thickets are so dense and exten- 
sive that part of a wild garden may be transformed into an impregnable fort 


285 Fig. 173. Alder swamp in Agri-chai valley. Biotope of jungle cat. Azeitaidzhan. 

December, 1957. Photograph by N.N. Rukovskii. 

(E.P. Spangenberg). When living close to man, the cat often intrudes into 
villages and steals domestic fowl. This cat has been found several times 
in barns, around dwellings, and other places. Jungle cats have sometimes 
been seen in the Trans- Caucasus in places that are pure steppe, several 
kilometers away from water sources (Smirnov, 1922). 

In Middle Asia this cat also inhabits reed thickets and tugais. How- 
ever, in Tadzhikistan in March and April, when desert hares [Lepus tolai] 
and pheasants migrate from the tugais into the adjoining desert to fatten 
up, the predator follows them there (Chemyshev, 1958). In the Amu-Darya 
delta it lives in extensive reed thickets, often flooded in summer, and is 
frequently sighted in bogs and breaks among the reeds a few kilometers 
286 from the banks, or on the shore (A.A. Sludskii, V.S. Pokrovskii). In Middle 
Asia this animal occurs in newly irrigated sections of the desert and inhabits 
thickets along the banks of canals and major irrigation ditches, which are 
reminiscent of miniature tugais. Pheasants, gerbils, hares, complex-toothed 
rats, and domestic mice share these habitats with such predators as jackal, 
steppe cat, and jungle cat. 

On the other hand, where the tugais have vanished the jungle cat has 


286 Fig. 174. Unfrozen opening on a river where tlie jungle cat hunt.s for wintering 

duciis and picks up injured ones. Severe, abundantly snowy winter, 1949/1950, 
in Trans-Caucasus. Kyurdamir, Azerbaidzhan. Photograph by S.S. Turov. 

become rare. For example, at places in Tadzhikistan where cultivated land 
has largely replaced tugais, this cat has become a rare animal. In the Gissar 
valley it survived only in infrequently occurring "islets" of reed thickets 
in the area of Gissar (Chemyshev, 1958). As a result of the destmction of 
its biotopes in recent years, the distribution and population of this cat in 
Azerbaidzhan has decreased rapidly (Aliev and Nasibov, 1966).^*^ 

As an inhabitant of lowlands, the jungle cat ascends only up to 600-970 
m above sea level in the Caucasus and Trans-Caucasus (Dinnik, 1914; 
Vereshchagin, 1947; Dal', 1954), to 900-1,000 m at Dushanbe in Tadzhikis- 
tan (Chemyshev, 1948), 1,800 m in Iran, and even 2,400 m above sea level 
in the Himalayas (Blanford, 1888-1891). 

Specialized for predation on animals closely associated with unfrozen 
water sources or those frozen only for a brief duration and not annually, 
the chaus finds optimum living conditions only in the far south of the Soviet 

-^The reports of G. Radde, A. Walter, [and W. Blasius]* (1889) that in the southern 
regions of Turkmenia, altogether devoid of shrubbery thickets, jungle cats were present and 
inhabited crevices and small caves in gorges on the banks of upper Murgab requires 
confinnation. These investigators report finds of jungle cat in such an environment on April 
8 to 20, 1887 at Takhta-Bazar. 

*Omitted in Russian original — Sci. Ed. 


Union. The northern Hmit of the massive wintering of waterfowl in the Soviet 
Union roughly coincides with the January isotherm of 2°. Evidently this 
isotherm also serves as the northern boundary of optimum conditions for 
the cat. It is not well adapted to subzero [Celsius] temperatures and snow 
cover. Of all our wild cats, the chaus has the shortest, coarsest, and sparsest 
hair coat, which poorly protects the animal from cold. Hence, in the northern 
part of the range during unusually severe and abundantly snowy winters 
this cat perishes from cold and hunger. In the severe winter of 1941/1942 
in Astrakhan preserve one cat came to a stable on February 1 and three dead 
ones were found in the Terkhizbenka River. The animals were emaciated 
and their intestines empty (Yu.A. Isakov). The absence or extreme rarity 
287 of this animal in Kazakhstan and the southeastern European sector, in spite 
of extensive reed thickets and swamps, is explained by the above ecological 
features. In addition,^now hinders cats venturing high into the mountains. 
Food. On the western shore of the Caspian Sea this cat feeds on water 
voles [Arricala terrestris], small mouse-like rodents, European hare [Lepus 
europaeus], ground squirrels [Spennophilus], wild boar (small sucklings), 
iasectivores (shrews), and in recent years the acclimatized nutria. It attacks 
ducks of various species, coots, moorhens, pheasants, francolins, gray part- 
ridges, and smaller birds; it also destroys fledglings and eggs. It eats turtles, 
snakes, lizards, and fish (mainly sazan). When living close to villages, it 
often steals chickens, ducks, and geese. It eats fresh carrion. Its main prey 
year-round are rodents and waterfowl (ducks and coots), especially in winter, 
followed by pheasants, francolins, and fish. In several regions the cat lives 
almost excliLsively on waterfowl. How diverse its prey is can be judged from 
the following examples. In the stomach of a chaus caught on the lower Terek 
(June) the remains of ten ground squirrels were found, and in the stomach 
of another caught in winter in Dagestan were some shrews swallowed whole 
(Heptner and Formozov, 1941). The stomach of a large male from Kyurdamir 
(Azerbaidzhan; June) contained two water voles, the scales of a sazan, and 
about 800 g of the meat of a small wild boar; the stomach contents of a 
cat from Kyzyl-Agach preserve (August) showed one water snake [Natrix 
tessellata or sipedoii], a pochard [Athaya ferina], and three house mice; the 
stomach of another caught in December contained a pochard; and that of 
a cat from Karayaz (Georgia; February) had one blackbird and two common 
voles (Vereshchagin, 1942). A marsh turtle was also found in the stomach 
of one cat. 

Jungle cats often hunt for nutria; for example in Karayaz (Georgia) they 
and jackals act as the chief agents limiting the nutria population. Even from 
farms fenced with a wire net to a height of 2.5 m, these cats stole 147 young 
and adult nutrias during the three winter months of 1947 (Vereshchagin, 
1950). A particularly large number of nutrias were killed by cats in the 


Fig. 175. Reed floodplains in "'Tigrovaya Balka"' piesei-ve. Habitat of jungle cat. 

Tiger, wild boar, Bukhara deer, and pheasants are also found here. Lower Vaklish, 

Tadzhikistan. October, 1960. Photograph by A.A. Sludskii. 

unusually severe winters of 1948/1949 and 1949/1950. In these winters, 
because water sources froze over, the cats caught nutrias hiding in thickets 
or running away over ice. The predator evidently catches mainly young 
nutrias and rarely attacks adults. A cat was once observed attacking a large 
nutria, which made no attempt to escape from its enemy, nor to dive into 
the water, but protected itself, and sometimes itself attacked. After 10 to 
15 minutes, the predator was compelled to withdraw, and after meowing 
swam away into the reed thickets (Pavlov, 1953). 

At such times [severe winters] cats kill many waterfowl weakened by 
hunger, the more so because they are concentrated in large numbers in a 
few frozen* stretches (E.P. Spangenberg). In the region of Kyzyl-Agach 
Bay in the severe and abundantly snowy winter of 1956/1957, little bustards 
[TetrcLx tetrax] and waterfowl wintering there had become greatly enfeebled 
and thus fell easy prey not only to foxes and jackals, but also to jungle 
cats (T.A. Adol'f). 

'Unfrozen? — Sci. Ekl. 


288 Fig. 176. Habitat of tlie jungle cat in "Tigrovaya Balka" preserve. Lower Vaklish, 

Tadzhikistan. April, 1966. Photograph by G.N. Sapozhnikov. 

While hunting for waterfowl, water voles, and nutrias, cats will often 
swim from bog to bog, sometimes across long stretches, not only in summer 
but even in winter. In hunting, upon reaching an opening among reeds, the 
cat simply lies in wait for any prey; noticing one nearby, frequently jumping 
directly on a swimming duck or nutria in the water (Kashkarov, 1932; M.P. 
Pavlov). In January, 1949, along a backwater of Lake Tomkin (Tadzhikistan), 
a chaus jumped from the bank onto ducks swimming in an opening in the 
ice and caught one of them (Chemyshev, 1958). 

In the summer of 1951, near Kyurdamir (Azerbaidzhan), as a result of 
intease drought many lakes dried up considerably. The water receded from 
the shore almost a kilometer, exposing extensive mud-flats; holes with water 
and fish in them remained here and there. In the drying lake from dawn 
to 7:00 to 8:00 a.m. cats hunted regularly, catching fish in these pits. Usually 
the predator covered the mud-flat between the bank and the hole in long 
leaps, and on seizing a fish, and quickly ran back into the reeds. The cat 
pounced in a long leap on sazans weighing 200 to 500 g moving in the 
shallow water, snatched one of the fish, and retreated rapidly to cover. 
Cats also preyed on fish at the end of the day, often even until sunset. In 


this same area and year they also stole water voles from traps (Yu.A. 

In regions where autumn and winter hunting for waterfowl is well 
organized, wounded birds and those weakened during overwintering often 

289 constitute the main quarry of the cat. In southwestern Turkmenia the main 

290 Table 12. Results of analysis of feces and stomach contents of jungle cats in Middle Asia 

(percentage of total number of observations) 

Region, season, and material 

Right tributaries Floodplains of Lower 

of upper upper Amu- Amu-Darya' 

Amu-Darya Darj'a, Aral- 

(Tadzhikistan)^ Paigambar Island^ 


— Summer Winter — 

100 samples 

33 samples 

(including 33 



(including six 


samples samples 










Desert hare [Lepus tolai 











Small jerboa [Allactaga 






House mouse [Miis musculus] 





Complex-toothed-rat [Nesokia] 





Midday gerbil [M. meridiantis] 





Red-tailed gerbil [M. eiythroiinis] 





Crested gerbil [Л/. tamariscinus] 





Great gerbil [Rhombomys] 





Mole vole [Ellobius] 





Trans-Caspian vole 





[M. transcaspicus] 

Muskrat [Ondatra] 





Nutria [Myocastor] 





Rodents, not further identified 





Boar (suckling) 





Jackal [Canis aureus] 










Pheasant [Phasianus] 










Pochard [Athaya ferina] 





Coot [Fulica atra] 







Table 12 (Contd.) 

Region, season, and material 

Right tributaries Floodplains of Lower 

of upper upper Amu- Amu-Dar>'a^ 

Amu-Darya Darya, Aral- 

(Tadzhikistan)^ Paigambar Island^ 

Summer Winter 

100 samples 

33 samples 

(including 33 



(including six 





Cormorant [Phalacrocorax] 





Black-necked sheldrake 





[Red-headed pochard, Netta 


Harrier [Circus] 





Birds, not further identified 





Pheasant eggs 





Reptiles and amphibians: 





Lake frog [Rana ridibunda] 





Steppe agama 





[A. sangiiinolenta] 

Box lizard [Ej-emias] 





Bluntnosed viper 





[V. labetina] 

Desert tortoise 





[Testiido horsfieldi] 

Eggs of turtle 








































Plant food: 





Oleaster nuts 





Green stalks of reeds 





Green stalks of sugar cane 




^V.L Chemyshev (1958). 
^G.I. Ishunin (1965). 
^A.M. Allayarov (1964). 


Fig. 177. Tugai along a dried up tributary. Habitat of jungle cat in "Tigrovaya 

Balka" preseive on Lower Vakhsh. Jackal, tiger, and pheasant also live here. 

October, 1960. Photograph by A.A. Sludskii. 

prey are rodents and birds, especially overwintering waterfowl, which are 
caught during the night, and francolins. Concerning food of the jungle cat 
in Middle Asia, see Table 12. 

It can be seen from Table 12 that in Tadzhikistan and Uzbekistan jungle 
cats feed mainly on desert hares [Lepiis tolai tibetaims], rodents (63-92% 
of samples), and birds (12-44%). Jackal remains in feces are evidently 
explained by the consumption of carrion. There is a definite place for plants 
in the diet of cats, especially oleaster fruits (Eleagmis angusti folia), the 
nuts of which are regularly encountered in feces. The food regime varies 
very little in different seasons but hares, complex-toothed rats, and pheasants 
predominate in the summer diet. In the Amu-Darya delta, in addition to 
birds and complex-toothed rats, the cat hunts acclimatized muskrats by 
watching at its dwelling or digging up the house; it destroys the constnictioas 
of this rodent more vigorously than does the jackal. In the muskrat harvest 
season cats approach very near to trapping parties and pick up discarded 
muskrat carcasses (A.A. Sludskii and V.S. Pokrovskii). 


In Afghanistan the stomachs of three chaus contained the remains of 19 
house mice, 3 complex-toothed rats, and 9 lake (?) frogs (Neithammer, 1966). 

The stomachs of cats have contained up to 1,200 g of food. In zoological 
gardens they are fed 720 g of meat with bones daily in summer, and 870 g 
in autumn and winter (Obukhova and Shakhnazarov, 1949). 

Home range. The size of individual home ranges are not known. Judging 
from tracks, they cover in one night no more than 3-5 km (Chemyshev, 
1958) or according to other data 5-6 km (Allayarov, 1964). 

Burrows and shelters. On the west bank of the Caspian Sea dens for 
kittens are established in openings among the reeds, on dry islets among 
thickets of blackberry and clematis, and in thorny green hedges. In the 
Lenkoran lowlands the cat often lives in deserted burrows of badgers, foxes, 
and porcupines. It lives in burrows in winter also. Statements to the effect 
that this cat digs its own burrows are unconfirmed. 

On June 26, 1925 in the vicinity of the city of Makhachkala a nest 
was found among reeds under a dense oleaster bush. The nearest reeds were 
pressed to the ground and served as the foundation of the nest. The [nest] 
platform itself consisted of short grasses and wool. The diameter of the 

Fig. 178. Thicket of giant plume grass and tamarisk in "Tigrovaya Balka"" preserve. 
Biotope of jungle cat. Tiger, Bukhara deer, wild boar, jackal, and pheasants also live 
here. Lower Vakhsh, Tadzhikistan. October, 1960. Photograph by A.A. Sludskii. 


foundation of the nest was 100 cm, and of the platform, about 50 cm; the 
thickness of the nest was 30-35 cm. The nest had been buih on a section 
of ground which was dry at that time of the year. Kittens with their eyes 
still closed were found inside the nest. A second nest with kittens was found 
on June 5, 1926 near Kizlyar. It was located on a dense reedy islet (area 
about a hectare) roughly 400 m from a small creek. The foundation of the 
nest consisted of reed stalks and the platform of soft grasses and wool. 
The overall diameter of the nest was 80 cm and of the platform 40-45 cm. 
Both of these two nests were well concealed (Yu.G. Heptner and Ognev, 
1935). In southem Turkmenia newborn kittens have been found lying directly 
on sand. Other cases of kittens found directly on the ground or under bushes 
are also known. 

In Tadzhikistan jungle cats set up nests in fortlike dense tugais or on 
fallen reed stalks in plume grass thickets. On these stalks, the cat trains 
dry leaves of reeds and other plants, forming a nest 1.5 to 2.0 m in diameter, 
covered from all sides with reeds. All three of the nests discovered were 
so constructed. Instances of cats inhabiting burrows in this area have not 
been recorded (V.I. Chernyshev, 1948). In Uzbekistan the jungle cat sets 
up its den in a similar manner in dense reed thickets. In the lower Zeravshan 
a nest 80 cm in diameter was found among fallen reeds and cattails of the 
previous years. The platform was covered with reed leaves and wool. From 
the platform a 2.0 m-long gallery imperceptibly opened into a space [in 
the reeds]. A second nest in the Amu-Darya deUa was on fallen reeds growing 
in shallow water 10 cm deep 80 cm from the surface (Allayarov, 1964). 
Near Samarkand, in the Zeravshan valley, a cat occupied the burrow of 
a porcupine at a distance from the irrigated land. Here there are neither 
tugais nor dense growths (V.G. Heptner). 

Daily activity and behavior. The chaus has essentially a nocturnal way 
of life, setting out to hunt at twilight, but in the warm season of the year 
it is often active in the morning and during the day. Outside the breeding 
period in the Trans-CaucasiLS it spends much of the day hiding in openings 
among reeds, thickets of thorny bushes, rock crevices, and from time to 
time in the hollows of trees (Dinnik, 1914); it also rests in burrows. 
292 The chaus is not timid and seldom wary. It falls into traps comparatively 

easily. It is strong and malicious; rare is the dog which can tackle one by 

While hunting on land the cat moves stealthily, without noise, close 
to the ground. It stops from time to time, listens, and sometimes raises its 
head as though scanning the distance. In stalking a quarry it almost creeps 
along the ground, hunches its shoulders tightly, pricks up its ears, spas- 
modically switches its tail, sometimes freezes in place, later creeps forward 
again, and finally makes a decisive leap. Its leaps are not long. If it misses 


on the first pounce, it makes one or two more leaps, after which the prey 
is not pursued. While hunting it is guided not only by audition and vision, 
but also by olfaction. Repeatedly cats have been observed to sniff constantly 
as they proceed along a path. Sometimes it bypasses well-hidden traps set 
in the trail. It often climbs trees; on one occasion a suckling [pig] killed 
and hung on a tree by hunters was partly devoured by a cat (Dinnik, 1914). 
Sometimes the cat lies in wait for its prey for quite some time, remaining 
concealed close to the trail in openings in reeds, and at other such sites. 
An observer once happened to see one jump from behind a bush on a desert 
hare running past and quickly dispatch it. 

Jungle cats usually hunt in the early morning and evening hours and 
rest at the end of the day. They also hunt in the first half of the night and 
have quite often been encountered searching for prey even during the day. 
Injured ducks in particular are frequently hunted during the day. In areas 
where hunting of game is common (Trans-Caucasus), cats regularly gather 
from the immediate vicinity. They are not afraid of gunshots, remain close 
to hunters, and quickly seize undetected wounded birds (E.P. Spangenberg). 
Magpies interfere in day hunts of jungle cats by following the predator 
and emitting loud alarm calls. 

Even when caught as a kitten, a jungle cat is quite difficult to tame. 
One animal, taken as a kitten only 30-40 days old, remained vicious the 
entire time. No one could extend a hand to it; it growled, attacked with 
its claws, and inflicted injuries even on a hand covered with a glove. This 
cat adapted to a diurnal mode of life but was especially active in the evening 
under artificial lighting. It was trained to jump from one stool to another 
and to walk a thin pole. While performing these exercises it issued loud 
calls and threatened the trainer constantly with its claws; it never attempted 
to bite him, however (N.N. Rukovskii). 

Seasonal migrations and transgressions. These aspects are not known. 

Reproduction. In Dagestan and the Trans-Caucasus, judging from the 
period of appearance of kittens, estrus would seem to be greatly prolonged, 
extending from the end of January through the first half of April inclusive. 
Hence statements (Vereshchagin, 1942) that mating in the Caucasus occurs 
in March and in the eastern Tfans-Caucasus in February (old calendar; 
Satunin, 1915) are only partly true. In southern Turkmenia mating occurs 
in January to early February. In southwestern Tadzhikistan the testes greatly 
increase in weight by January and become particularly large in February- 
March. The testes of a cat caught in October weighed 1.03 g while those 
of another animal caught in February weighed 4.62 g. 

Active spermatogenesis occurs in February and the first half of March 
but, considering the period of the appearance of kittens, one could assume 
that it occurs even in January. In the middle of March spermatogenesis slows 


down but spermatozoa are preserved in the epididymis until mid-May (two 
cases). Based on the condition of the testes, оссттепсе of pregnant females, 
and periods of birth of kittens, one may assume therefore that the period 
of estrus usually extends from January to March in the region under con- 

In warm winters estrus commences almost one month earlier than in 
severe winters. In "Tigrovaya Balka" preserve on the lower Vakhsh the 
earliest mating of these cats was observed on January 10, 1948 and the 
last on February 15, 1950. It is possible Ihat in spring some individual females 
mate even later. In 1950 two adult females, not yet pregnant, were caught 
on March 25 and 30 and another nonpregnant female even on April 10 (V.I. 
Chemyshev). During the period of estrus, after sunset and throughout the 
night, the shrieks of males are heard, which differ from the normal howls 
of domestic cats in a more prolonged "gurgle". Violent combats occur 

293 between males in rut as evidenced by traces of blood and tufts of hair. The 
duration of gestarion is not precisely known but evidently covers a period 
of about 60 days. 

In Dagestan kittens are seen from the end of March to early June. The 
earliest kittens have been discovered in the Kyzlyar region was the last few 
days of March, 1926. Near Makhachkala three kittens with their eyes still 
closed were discovered on June 20, 1925. Two fairly grown kittens were 
found near Kyzlyar on June 5, 1926 together with a lactating cat; another 
very large kitten was found here in the first half of July, 1925 (Yu.G. Heptner, 
Heptner and Formozov, 1941). In the Trans-Caucasus (Lenkoran region) 
on June 14, 1933, a litter of three kittens with their eyes open but still weak 
in their legs was found (E.P. Spangenberg), and another three in the same 
condition happened on May 25, 1951 at Kyurdamir (Azerbaidzhan). Two 
females caught at Kyurdamir on June 7 and 10 were found to be lactating 
and had considerable milk in their glands; four placental scars were found 
in the uterus of one (Yu.A. Gerasimov). 

In the Caucasus young are seen in early May (Vereshchagin, 1942). 
In southern Turkmenia at Saryyaza on March 29, 1887 (old calendar) five 
newborn kittens were found (Radde, 1889). Kittens with a body length of 
about 20 cm were found on April 5, 1901 (old calendar) in the Tedzhen 
valley (Zoological Museum, Academy of Sciences). In southwestern Tad- 
zhikistan the first pregnant female was caught on February 13, 1948, and 

294 contained three fetuses weighing 13.7 to 14.2 g. A second cat caught on 
February 21, 1950 was found to be in estrus. Of two females caught on 
March 3, 1949, one had four fetuses aged about 30 days and the other three 
aged 20 to 25 days. A cat caught on March 21, 1950 had only just mated; 
three софога lutea were found in her ovaries. 

The earliest date of all births recorded was March 31, 1949; this parti- 


293 Fig. 179. Impenetrable stronghold of oleaster {Eleagnus), plume grass, reeds, and 

liana. Residence of jungle cats in Pyandzh valley. Tadzhikistan. April, 1966. 
Photograph by G.N. Sapozhnikov. 

cular female produced four kittens. A female caught on April 18, 1950 was 
already lactating. The last of the births recorded took place on May 10, 
1947 (Tadzhikistan, Parkhar region) (Chernyshev, 1958). Judging from the 
periods of estrus and sightings of pregnant females, kittens in the Parkhar 
region (Tadzhikistan) may evidently be expected anytime between mid- 
March to mid-May, possibly even later. 

The litter consists of three to five young, usually only three (14 obser- 


vations). Among cats breeding in Alma-Ata Zoological Garden there were 
usually three kittens per litter; four were found only once. References to 
the occurrence of up to 10 kittens in a litter are apparently erroneous 
(Kashkarov, 1932; Sultanov, 1939; Naumov and Lavrov, 1941). 

The well-known report of Blanford (1888-1891), quoted by later 
investigators, mentions that in India this cat annually produces two litters 
of three to four kittens each. This cat sometimes produces two litters in 
a year in the Soviet Union also. Thus in Ismaillinsk region of Azerbaidzhan 
pregnant jungle cats were encountered in summer also. At the end of August, 
1952, a nest deep under the roots of a tree contained kittens about 30 to 
40 days old (their eyes were open and their teeth empted) (N.N. Rukovskii). 
In the Zoological Museum, Academy of Sciences, there is a kitten weighing 
one- fourth the weight of an adult with the label "19 October, 1936, western 
Kopet-Dag, Chandyr River". This cat must have been born in August. A 
pair of jungle cats from southern Turkmenia littered twice in a season in 
Alma-Ata Zoological Garden, i.e., in March-April and in August; in 1957 
they even produced three litters in a single season (the female did not nurse 
the young). In Uzbekistan in the Angren valley kittens have been seen at 
the end of April and in early August (Lustin and Putsatov, 1957). 

In Tadzhikistan a high percentage of barren females has been noted 
among female jungle cats. Of the 11 aduU females caught during the breeding 
season, only 7 (63%) were pregnant and 4 (37%) barren. The reasons for 
this phenomenon have not been established (Chernyshev, 1958). 

Growth, development, and molt. Kittens are born blind, with their ears 
closed, and helpless. They are covered with a dense furry coat of hair. The 
general color shade of the juvenile coat on the head and back is a light brown, 
gradually fading toward the flanks. The underside of the body is light olive. 
On the forepaws there are four faint transverse bands, and on the rear paws, 
six. Faintly visible dark-colored spots occur on the flanks and underside 
of the body. In the rear of the back there are four longitudinal dark bands. 
Longitudinal bands also occur on the crown and neck. The tail is the same 
color as the back. Among kittens one-fourth the size of adults, bands are 
present on the rear paws and flanks. The weight of four two-day-old kittens 
born on March 31, 1949, was 45, 50, 43 and 55 g. 

The kittens gain sight on the tenth to the twelfth day. lactation continues 
for about two months. In southwestem Tadzhikistan the cessation of lactation 
in different years varied between June 10 and July 20 (Chernyshev, 1958). 
Milk teeth were shed by a young captive cat and replaced by permanent 
ones at the age of five months (N.N. Rukovskii). 

The litter of this species breaks up early, at the end of summer, and 
hence the animals are invariably seen singly in autumn and winter. By 
October-November the young sometimes attain one-half the size of adults 


and weigh at that time 5.0 to 6.0 kg. In winter they grow very slowly and 
hence young animals in spring weigh the same as in [the previous] autumn. 
In southwestern Tadzhikistan young females weighed 2,400 g on November 
12, 1949; 2,700 g on March 30, 1950; and 2,618 on April 1, 1950 (the weight 
of the largest adult female was 7,500 g). A young male caught on March 
5, 1947 weighed 5,000 g (the adult weighs about 8,000 g, and even up 
to 12,000 g) (Chemyshev, 1948). A tame chaus aged 7.5 to 8.0 months 
weighed 7.0 kg and was not obese (N.N. Rukovskii). 
295 The jungle cat molts twice a year. In the Trans-Caucasus spring molt 

commences in February and ends by mid- April. Cats caught in June sport 
a summer coat and a clean light-colored inner skin surface. Autumn molt 
is more protracted. It commences in September and ends in the first few 
days of November. Only after this period does the skin become completely 
renewed. Reference to molt occurring in May in the Caucasus is errone- 
ous (Veres hchagin, 1942). In southwestern Tadzhikistan in warm winters 
jungle cats begin to molt in the first few days of February. The limbs 
and heads of specimens caught in the middle and end of February were 
completely mohing. By the end of April all animals have completed the 

In the cold winter of 1949/1950, when very low temperatures prevailed 
from December to beyond the first half of February, molt was delayed. Cats 
caught in early and middle March still bore a winter coat with faint traces 
of molt. Nevertheless, by early May a summer coat was seen in all animals. 
Thus, in this cold winter the commencement of molt was delayed by one 
month. In autumn the shedding of summer hair (guard hair) and the growth 
of underfur were seen from mid-October on. Molt was completed by the 
end of November to mid-December. The winter coat is more grayish in 
hue while the summer coat is more sandy, blending well with the surrounding 
dry vegetation (Chernyshev, 1958). 

Enemies, diseases, parasites, mortality, competitors, and population 
dynamics. Enemies and diseases are not well known. Kittens caught in 
Ismaillinsk region (Azerbaidzhan) were infected with coccidians (N.N. 

In Georgia the following helminths were found in chaus: Hydatigera 
taeniaeformis,Mesocestoides lineatus, Toxocara mystax, CapiUaria feUscati, 
Thominx aerophilus, and Ancylostoma caninum (Rodonaya, 1951). In 
Azerbaidzhan seven species of parasitic worms have been found: Diphyllobo- 
thrium erinacei, Hydatigera krepkogorskii, H. taeniaeformis, Mesocestoides 
lineatus, Troglostrongylus assadovi, Toxocara mystax, and Petrowospirara 
petrowi (Sadykhov, 1955). In southwestern Tadzhikistan 10 species of 
parasitic worms and one parasitic larva have been detected (Chemyshev, 
1953) (Table 13). 


Table 13. Helminth parasites in jungle cats of southwest Tadzhikistan 
(upper Amu-Daryii; Chernyshev, 1953) 

Helminth or its lai-va 

Number of Degree of Number of Location of 
infected infection parasites parasite 
animals (percen- 

Diphyllobothnum maiisoni 




Small intestine 

Sporganum mansoni 




Body cavity, 
subcutaneous tissue 

Hydatigera taeniaeformis 




Small intestine 

Mesocestoides Uneatus 




Large intestine 

Toxocara mystax 




Gullet, abdomen, 
and intestine 

Uncinaria stenocepliala 




Large intestine 

Gnatostoma spinlgenim 




Abdominal walls 
and cavity 

Dirofilaria immitis 





Dirofilaria repens 




tissue, capsule, 

Dracimcuhis medinesis 




Subcutaneous tissue 

Pity Salop tera praepii tialis 





In Uzbekistan the bronchi of one of eight cats dissected were infected 
with the nematode Рга^/о5Гга/г^у/м5 badan'mi (Muminov, 1964). The number 
of helminths in a kidney, usually the right one, can be so high that its weight 
may exceed the weight of the other by twofold (Reimov and Nuratdinov, 
1970; Amu-Darya delta). In southwestern Tadzhikistan, the ticks Rhipice- 
phaliis turanicus, R. sanguines, another nonidentifiable Rhipicephalus species, 
296 and Hyallomma anatoUcum were found in these cats. Maximum infestation 
by ticks was observed from May through September (V.I. Chernyshev). 

In southwestern Tadzhikistan the fleas Pulex irritans and Xenopsylla 
nesokiae were found in two animals. The latter flea is a parasite characteristic 
of the complex-toothed rat. 

The competitors of jungle cat are jackals, forest and steppe cats, foxes, 
and predatory birds, especially the marsh harrier [Circus aeniginosus]. The 
main competitors are jackals and forest cats, which feed on almost the same 
animals. Their importance as competitors can be judged from the following: 
in the Trans-Caucasus, near Shil'yan, 290 jackals and 68 jungle cats were 
caught around a group of lakes in the winter of 1947/1948. Usually jungle 
and forest cats do not live in the same [geographic] place or biotope. If, 
however, they occur in the same region, the one species is always the only 


one. It is significant that jungle cats are absent on the Black Sea coast of 
the Caucasus and in the swamps of the Kuban, where forest cats are abundant 
or common. 

Judging from the record of the tanned skins of chaus in the southern 
part of its range, its population fluctuates from year to year, but not much, 
deviating from the multiyear mean by not more than 30%. Factors respoasible 
for fluctuatioas in population numbers have not been studied. In the northern 
part of the range fluctuations in population numbers are evidently due to 
extreme cold and abundantly snowy winters, when all water sources freeze 
and deep snow cover prevails for a prolonged period. In the southem part 
of the range, however, these very same conditions are distinctly favorable 
for the cat; in such winters it enjoys an abundance of food in the form of 
aquatic and other birds weakened by lack of food. In some years, however, 
the southem population may undergo a reduction due to epizootic diseases 
(carnivore distemper, pasteurellosis, paratyphoid, rabies, and others) (also 
see ''Population"). 

Field characteristics. A large cat (weighing up to 12 kg), standing tall 
on its legs, with a monochromatic reddish-brown coat. The tail is rather short 
and thin and the ears erect with tassels. Usually, it is seen running in an 
unhurried, slow gallop. While running, it sways slightly from side to side. 
It is often encountered near water, in openings in reeds, in kupak*, or while 
swimming. Its tracks are often seen on sandbars, spits, and silty banks and 
are much larger than the prints of domestic, steppe and forest cats. The 
imprint of the paw of a jungle cat is 5.0 cm x 6.0 cm in size. Moreover, 
the digital impressions are widely separated, while in other cats they are 
compact. The impression of the hind sole is trapezoidal and not rounded 
as in the forest cat (Fig. 180). Tracks extend in the form of zigzag line. 
The pace length is 29 to 32 cm. 

The meow of a jungle cat is similar to that of domestic cat but deeper. 
The voice is heard rather frequently at night, especially at the end of winter 
and in spring. (A.S.) 

Practical Significance 

Jungle cats are fur-bearing animals but their pelt are of little value. The All- 
Union standards include their pelts in the group "wildcats," in which they 
are distinguished as "jungle cats {sabanchiy\ 
297 Commercial hunting of the jungle cat was practiced even in the 

eighteenth century. P.I. Rychkov (1762) wrote: "Wildcats were present 

*Floating reed mats — Sci. Ed. 


296 Fig. 180. Tracks of a jungle cat on silt. "Tigrovaya Balka" preserve at Vakhsh. 

Tadzhikistan. October, 1960. Photograph by A.A. Sludskii. 

among Kirgizian hordes on the steppes, but few brought in for sale by the 
Kirgiz-Kaisak in Orenburg and Troitsk Fort. They were far bigger than 
domestic cats and resembled lynx, though smaller, and the fur was poorer. 
A single animal sold for 16 to 20 kopecks." Judging from the description 
of the skin these were jungle cats. 

At present, in fur-tanning factories this cat is of some importance only 
in Azerbaidzhan, where 2,966 to 4,300 or an average of 3,500 skins were 
tanned annually from 1931 through 1940 (these also include pelts of forest 
wildcat) (Veres hchagin, 1942). In Tadzhikistan in 1946 about 300 skias were 
prepared (Chernyshev, 1948). In Turkmenia the tanning of pelts of jungle 
cat has fallen considerably in recent years, mainly because of the absence 
of commercial hunting. Here 91 animals were caught in 1954, 109 in 1955, 
187 in 1956, 172 in 1957, and 166 in 1958. There is no information for 
the rest of the Union Republics since jungle cat is reckoned as "wildcat" 
by fur-tanning factories. 

This cat everywhere is a serious pest of the hunting economy. All 
investigators regard it as the main enemy of pheasants, francolins, and 


waterfowl. At several places, although the francolin was protected from 
hunting, its population did not rise due to its destruction by the jungle cat 
(Satunin, 1915). Like the wolf, this predator greatly damages the nutria 
farming of a semiwild type. Next to and within the precincts of a nutria 
farni about 200 cats were caught in traps over a period of 14 years. In 
Kyurdamir (Azerbaidzhan) since 1946, a bounty of 5 rubles is paid by nutria 
farms for the skin of a cat. At places, especially in the Amu-Darya delta, 
the cat adversely affects the muskrat economy. In some regions this cat 
is also a pest of poultry farms. 

No special hunting is organized to catch this cat. It is caught incident- 
ally during hunts for pheasants and pigs; it also falls into traps set for 
fox or jackal. It can be caught comparatively easily in a trap placed in 
water at a shallow place with a live fish fastened to a rod run through 
its gills. The splashing of the stmggling fish attracts the attention of the 
cat (Yu.A. Gerasimov). Pelts are used in making ladies' fur coats or imitation 

In view of the harm inflicted by this predator, its capture is permitted 
everywhere year-round. In Azerbaidzhan a bounty of 5 to 12 rubles is paid 
for each skin brought in. 

In old literature there exist references that in ancient Egypt, judging 
from mummies found, the jungle cat was domesticated; but later it was 
not found in a domesticated state (Bogolyubskii, 1959). Ancient Egyptians 
successfully crossed domestic cats with chaus and mummies of such hybrids 
have purportedly been found. It has also been reported that hybrids were 
used in hunting waterfowl and for tracking wounded birds; some pictures 
have been preserved depicting various scenes of such hunts with hybrid 
cats (Bogdanov, 1913). These data have found no confirmation in recent 
literature (see the section on the origin of domestic cats under group libyca 
and "Systematic Position" of jungle cat). (A.S.) 


Felis (Felis) silvestris Schreber, 1777 

1777. Felis (Catiis) silvestris. Schreber. Sdugeth., vol. 3, p. 397. Germany 

or northern France. 

In the species F. (F.) silvestris are here included two groups of forms 

eariier coasidered independent but closely related species: F. (F.) silvestris 

Schreber, 1777 — forest cats of Europe, Asia Minor, and the Caucasus, and 

F. (F.) libyca Forster, 1780 — steppe (spotted, bay, and long-tailed) cats 

of Africa and Asia. 

298 The basis for this unification was indicated in the last century but 


formalized only in the present-day (Haltenorth, 1953). Both species are 
moфhologically very similar and there are no constant craniological 
differences between them in the various parts of their range (Pocock, 
1951).^ Differences between the two species are confined to coloration 
and some features of the pelage (general characteristics of fur and fluffi- 
ness of tail). Within the limits of each "species" geographic variation 
does occur but nonetheless the distinctive features of both are linked by 
intermediates. Both "species" and all their geographic forms are strictly 

All forms belonging to the European forest cat group (silvestris), like 
those of the steppe cat group (libycd), hybridize in nature with the domestic 
cat, which itself is a domesticated steppe ("bay") cat of northeastern Africa, 
F. s. libyca (see p. 452). In some regions of Europe it is believed that the 
entire population of wildcats has some, even significant, admixture of the 
blood of domestic cats, and that evidently the hybrids may be regarded as 
299 fully fertile (this has also been established experimentally). Domestic cats, 
especially in countries with snowfree winter or those with little snow, readily 
tum feral, and some populations asserted or suggested to be separate species 
or subspecies merely represent feral domestic cats {megalotis — Timor, 
agria — Crete, (iaemo /г— Caucasus, mediterranea — Italy, pulchella — 
Africa, syriaca — Syria, and issikkulensis — Semirech'e). 

The range of the species silvestris, as interpreted here, is very extensiye 
(Fig. 181). However, it is unified and quite natural. The range of this cat 
is quite similar, broadly speaking, to the range of other widely distributed 
cats of the Old World (lion, leopard, caracal, and cheetah). The range 
occupies Europe, including England in the west (absent in Ireland and never 
present), the BaUic region in the north, and up to the Dnepr and Don Rivers 
in the east; Africa except the dense humid forests of western Africa; the 
Near East, including Asia Minor, the Caucasus; northwestern India; Middle 
Asia and southern Kazakhstan, Dzhungaria, Kashgaria, Inner Mongolia, 
and southern parts of Mongolian People's Republic (reconstructed range). 

The entire mainland of Europe and England are inhabited by typical 
members of the silvestris group. The same is true of the Caucasus and Asia 
Minor, at least over much of it. Both in the Caucasus and in southern Asia 
Minor, the ranges of subspecies of the silvestris approach very close to, 
and adjoin at places, the ranges of subspecies of the libyca group. Transitional 
populations however are absent; and hence transitional animals are not 
known there. 

-^^Craniological differences detected between Turkestan steppe and European 
(Caucasian) forest cats (V.G. Heptner; see pp. 75 and below), not observed by earlier authors, 
could not be verified in all the geographic forms. 




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Fig. 181. Reconstructed species ranges of forest and steppe wildcat, Felis (Felis) 
silvestris Schreb. s. I . (schematic) (scale in km). Region of occurrence of European 
forest cat — subspecies group silvestris (in the northwest), and steppe cat — subspecies 
group libyca (rest of the range) separated by broken line. Mediterranean insular 
transitional forms are included in the European forest group. The northern boundary 
of the range in Kazakhstan is tentative (see Figs. 198 and 267). (V.G. Heptner). 

The Mediterranean island forms and in part those of northern Africa 
[jordansi, reyi, cretensis {agria), and sarda] represent intermediates. 
Whatever be the characteristics of these individual forms and their history 
(degree of admixture of blood of domestic cats, domestic cat simply turned 
feral, etc.), some of their features are a mixture of features of European forms 
of the silvestris group and of the African libyca group. In the nature of 


their fur and color and fluffiness of tail, they belong to the libyca type, 
but the skull shows features of the silvestris type.^ 

Even the diversity in their interpretation underscores their unique 
(intermediate) taxonomic position. At the beginning of the century (Miller, 
1912) they were considered completely independent species {sarda and 
agria-cretensls) but nowadays all have been combined into one subspecies 
(sarda); some taxonomists place the subspecies in species libyca (Pocock, 
1951; EUerman and Morrison-Scott, 1951 and 1966), while other maintain 
that they occupy an intermediate position between the "species" silvestris 
and libyca (Haltenorth, 1953 and 1957). This latter view is very widespread. 

The systematic relations of the two groups of forms viewed from the 
process of speciation are of extreme theoretical interest. They may be 
assigned an intermediate position between species and subspecies or the 
entire complex of forms considered "superspecies" or "macrospecies". 

A description of the species, including synonyms, has been given 
separately for the two "groups" of subspecies: European wild forest cats, 
F. (F.) s. — ' 'silvestris group"; and steppe wildcats, F. (F.) I. — ''libyca 
group". Only in the sections "Diagnosis" and "Systematic Position" have 
the two been presented together. (V.H.) 


Size small; like that of domestic cats or only slightly larger. 

Background color gray of different inteasities, being very light-colored 
300 in some forms with a fairly intense ocherous shade. Dark spots are present, 
or a dark belt and spots on the back, or a belt and transverse stripes on 
the trunk, or a belt without distinct spots or transverse bands. Dark 
longitudinal bands occur along the top of the head. The tail has dark spots 
or rings (transverse). The coat is thick and rich with dense underfur, or 
relatively short with slightly developed and close-fitting underfur. Tail furry, 
thick, and blunt, or thin and pointed at the end. 

Skull not elongate, and zygomatic arches wide-set. Palate not extended 
into inteфterygoid vacuity posteriorly (see Fig. 28) and inteфterygoid vacuity 
broad. Second premolar (first in row) generally present; antero-inter cusp 
of upper carnassial tooth well developed and its peak distinct but small and 
blunt. Ectotympanic chamber of auditory bulla poorly developed (see Fig. 
24). Maximum diameter of auditory opening not more than length of third 
(second in row) upper premolar and part of auditory bulla between auditory 

"^One of the characteristics of these forms, judged from the limited amount of material 
from Sardinia {sarda) available in the Zoological Museum, Moscow University, is the 
similarity in structure of auditory bulla to that oi silvestris forms, i.e., the small bulge in 
front of the ear canal in the ectotympanic part is absent (see Fig. 29). 


and Eustachian openings does not bulge (European wildcat group, silvestris); 
or maximum diameter of the external auditory meatus more than length 
of third upper molar and auditory bulla between auditory and Eustachian 
openings somewhat inflated and shifted forward slightly more than portion 
lying inward from Eustachian tube (steppe cat group, libyca; Fig. 29). (V.H.) 


1777. Felis (Catiis) silvestris. Schreber. Saugeth., vol. 3, p. 397. Germany 

or northern France. 
1777. Felis catus ferns. Erxleben. Syst. RegniAnim., vol. 1, p. 518. Southern 

1837. Fells cato ajfinis.'^ Hohenacher. Bull. Soc, Naturalistes Moscou, 

1, 7, 136. Elenendorf, Elisavetpol' governance (now Khallar, south 
of Kirovabad, Azerbaidzhan). 

1896. Catus ferox. Martorelli. АШ Soc. Ital. Sci. Nat. Milano, vol. 35, p. 
275. Thuringia. This name arose through some misunderstanding 
(described together with Catus ferus Brehm, 1869) but was never- 
theless used in the nomenclature for this form in the Russian fauna. 

1904. Felis daemon.^^ Satunin. Proc. Zool. Soc. London, p. 162, Elisavet- 
pol' (now Kirovabad, Azerbaidzhan). 

1905. Fells catus caucasicus. Satunin. Izv. Kavk. Muzeya, Mitt. Kauk. Mus., 

2, 2-4, 154. Borzhom. 

1916. Felis silvestris trapezia. Blackler. А/ш. Mag. Nat. Hist., vol. 18, p. 

73. Khotz near Trapezund [Trabzon], Asia Minor. 
1943. Felis silvestris euxina. Pocock. Лл/г. Mag. Nat. Hist., vol. 10, p. 701. 

Baspunar in Dabruja, Romania. (V.H.) 


Size small, but averaging larger than that of domestic cats. 

In general appearance the forest cat is very similar to the domestic 
cat — body long and stands on rather short legs, and tail comparatively long 
(Fig. 182); its length usually slightly exceeds one-half of body length 
(sometimes comprising 70% of it), or equal to it, and more rarely shorter. At 
the same time, partly due to the generally greater size, but mainly because 
301 of the much longer and richer coat, especially the winter coat, the forest 
cat appears far bigger, powerful, and more massive than the domestic cat. 

Head broad; jugal region with short, blunt maxillary portion seemingly 

-'*Black feral domestic cat or melanistic forest cat. 



Fig. 182. European wildcat. Berlin Zoological Garden. Photograph by G. Budikh. 

less extended — skull more spherical than in jungle and Amur cats. Ears 
moderate in length, quite wide-set, and without tufts at tip. Hair at tip 
nevertheless may be somewhat long — up to 3.0 to 4.0 mm. Ears in winter 
project out of fur. Side whiskers absent on cheeks. Vibrissae white; hairs 
on lips range from 7 to 16 on each side and reach 50 to 80 mm in length, 
eyelashes number 6 to 8 each and range from 50 to 60 mm in length; group 
of 3 to 6 vibrissae 30 to 40 mm in length present on inside of wrist 
(Haltenorth, 1957). Eyes large, pupil vertical, and iris yellowish-green. Bare 
region on nose in living animal flesh-colored (not black). 

Pelage throughout body fairly uniform in length (long hair absent on 
neck or back). Hair on tail very long and dense and hence tail appears furry 
and thick. At end of tail hair not shortened and tips not pointed but thick as 


though chopped (characteristic feature of forest wildcat). Length of guard 
hair in winter coat 70 to 72 mm, top hair 55 to 60 mm, and underfur 45 
to 55 mm; corresponding measurements in summer coat 50 to 67, 45 to 
60, and up to 53 mm. Thickness of top hair 90 to 100 microas and of underfur 
20 microns. On the average 10 to 20 guard hairs, 65 to 100 top hairs, and 
1,600 to 2,100 underfur hairs occur in an area 25 mm^ of the winter coat 
along the middle of the back; corresponding figures for the summer coat are 
6 to 9, 66 to 100, and 380 to 1,100 (central European form; Haltenorth, 1957) 
(see "Geographic Variation" for description of pelage of Caucasian cat). 
302 Main color shade of winter coat fairly light, gray, somewhat richer 

along the back and faded toward the undersides of the flanks.^ Ocherous 
bloom imperceptible or barely discemible only along underside of flanks. 
Very distinct, fairly narrow black band commences at the shoulders, runs 
along the back and usually terminates slightly short of tail base; however, 
it sometimes reaches the tail or even extends onto it as a narrower band 
along the dorsal surface. 

Color on flanks monochromatic; transverse stripes from dorsal band 
either absent or present only as hazy, indistinct shadows; they are rarely 
distinguishable as stripes and better identified as indistinct dark smudges; 
sometimes, especially on underside of flanks, they have a rusty tone. Fairly 
distinct but not very sharp transverse stripes, not more than five to seven, 
are a rare occurrence. Transverse bands on the legs are either vague or 
undeveloped and only occasionally fairly distinct. 

Undersurface of body covered with very light gray hair with a light 
ocherous tinge; dark spots absent or occasionally a few present. On the 
throat a white spot may be present (quite rarely) or three such spots occur — 
one on the throat, another between the forelegs, and a third in the rear 
(inguinal) region of the abdomen. Posteriorly the abdomen is generally a 
pure and quite bright ocherous color. 

Color of tail same as on the back; tip of tail pure black. Above tail 
tip usually two, more rarely three clear, black, transverse rings occur, which 
are narrower than the black tip and separated from it and from each other 
by light-colored sections; width of sections roughly equal to the width of 
the rings. Usually, in front of the black rings, two or three more indistinct, 
rather dull, sometimes barely perceptible dark-colored, most often brown 
rings occur. Sometimes along crest of tail there lies a narrow dark band, 
which is a continuation of the band along the spine that joins the rings on 
the tail. This pattern is found in cases where the number of rings is greater 

'^''Description based on a series of Caucasian skins available in the Zoological Museum, 
Moscow University. This description differs significantly from that given by S.I. Ognev 
(1935) since the latter had only limited material available to him and that, too, not wholly 
representative but with a relatively well -developed pattern. 











and their color brighter, and general color of the tail also more vivid. Main 
color of lower part of the tail slightly lighter than upper part. 

Upper surface of front paws monochromatic, grayish-yellow, and same 
color as foot but more distinctly rust-toned. On the lower surface of the 
foot, in the anterior part, a dirty blackish-brown section covers one-third 
or one-half the area of the paw (remainder dirty ocherous-gray) or sometimes 
the whole of it.^^ 

Dorsal surface of the neck and head same color as dorsal surface of 
the trimk, but around the eyes, lips, cheeks, and chin color lighter gray with 
a fair admixture of ochre. Hair on inner surface of ear yellowish-white and 
also forms a sort of border on outer surface. Outer surface of ears dark 
gray with a rusty or ocherous tone. Forehead and top of head bear pattern 
of four well-developed black or brownish-black bands; sometimes these 
bands split, on forehead, into small spots extending onto neck. In the shoulder 
region, in front of black dorsal band, usually two short, narrow, dark stripes 
present. From outer corner of the eye, under ear, and beyond onto side of 
neck, runs a dark narrow stripe; another similar stripe commences under 
the eye, slightly away from it, and also extends onto neck. 

Summer coat with fairly light, pure background color, without admixture 
of ocher or brown; it is even ashen-colored in some animals. Pattern on 
back and head well developed like pattern on tail, but pattern on flanks 
very indistinct and may be aUogether absent. 

There are no sex-related differences in color although possibly females 
may be somewhat more gray than males (Dinnik, 1914). Pelage of young 
303 animals at birth has a very light general background with very vivid velvety 
black spots. Bands and spots numerous, sharply defined, vivid, and cover 
greater area than general background color. Spots in adult animals possibly 
fainter than in old animals. 

Geographic variation in color is not pronounced. The intensity of 
general coat color and number of spots and stripes and their degree of 
vividness vary moderately. On the whole coloration conforms to Gloger's 
rule, with greater vividness and more prominent spottiness in forms from 
Scotland, and an intermediate coloration evident in central European 

Preanal glands present in both sexes; moderate-sized sweat and sebace- 
ous glands are located around the anal opening and larger-sized sebaceous 
glands and a few scent glands dispersed in the caudal region, extending 
along the entire tail on dorsal side. In males, preanal pockets are present in the 

^^The dark field, when it covers the entire undersurface of the paw, is usually considered 
(Suminskii, 1962; see later) due to an admixture of blood of domestic cat. Among Caucasian 
cats this occurs merely as an individual variation. 


303 Fig. 183. Skull of Caucasian forest wildcat, Felis {Felis) silvestris caiicasica 

Satunin. No. S 14230, collection of the Zoological Museum, Moscow University, 
old male. Near Tarsk station close to Ordzhonikidze. December 6, 1928. Sketch 
by N.N. Kondakov. 

form of two folds on each side of the anal opening with channels for 
304 discharging secretion. These pockets, like the glandular area along the dorsal 


surface of the tail, are activated with the onset of sexual maturation and 
play a significant role in reproduction and marking of territory (Haltenorth, 
1957; for domestic cat). 

Teats, four pairs — two thoracic and two abdominal. 

The skull of the forest wildcat (Fig. 183) differs significantly from that 
of the jungle cat in somewhat smaller size and weight (for the same age). 
It is significantly less extended and the zygomatic arches more wide-set 
not only medially (zygomatic width three-fourths of condylobasal length), 
but also anteriorly, where they are more steeply set in relation to the sagittal 
plane. General outline of skull also more spherical since the cranial region 
is not so constricted and extended posteriorly. Upper line of profile bulges 
while the large frontal area bulges only slightly, or is almost flat, or exhibits 
a very gentle and barely perceptible depression. 

Orbits relatively large, quite rounded, and set forward to a greater degree. 
Supraorbital processes project perceptibly downward and often are so well 
developed that they almost meet postorbital processes of jugal, or even fuse 
with them to close the orbits from behind. In front of orbits, maxillary (facial) 
section of skull highly compressed and jaws therefore parallel. Nasals 
generally terminate posteriorly quite bluntly, but in some individuals rather 
sharply; distance between anterior processes of frontals and upper ends of 
premaxillae greater. Lateral margins of nasals highly compressed medially 
and relatively broader anteriorly than in jungle cat. Sagittal crest in old 
animals well developed but only in posterior part of parietals; lambdoidal 
crests fully developed, and relatively stronger. 

Auditory bulla occupies roughly the same area as in jungle cat but 
smaller in size since less inflated, especially in ventral and anterior directions. 
Anterior margin of bulla does not reach level of glenoid fossa and postglenoid 
process. Maximum diameter of opening of auditory meatus less than length 
of upper third premolar (anterior to carnassial tooth) or equal to it. Bulla 
not inflated in front of opening of auditory meatus and chambers on either 
side of Eustachian opening roughly equal in size and lie at approximately 
the same level. -"^^ 

Palate not extended posteriorly into inteфterygoid vacuity; its posterior 
boundary either lies at level of posterior edge of molars or only slightly 
projects beyond that level. Inteфterygoid vacuity relatively narrower 
than in jungle cat and more uniform in width throughout its extension. 
Posterior edge of palate with small medial tapered projection. Palate 
itself relatively shorter than in jungle cat and somewhat relatively broader 
posteriorly. In conformity with more sharply sideward-set zygomatic 
processes of maxillae, tooth rows do not form a straight line from first 

•'-See notes about this characteristic under description of steppe cat. 


maxillary tooth to axis of carnassial as in jungle cat; instead this line is 
somewhat arced posteriorly. 

Dentition complete, i.e., second upper premolar (first in row of maxillary 
teeth) present and functional. However, this tooth is frequently absent. Thus, 
they were absent on both sides in 6 (35%) out of 17 males examined from 
the Caucasus. Among all but 2 skulls of 23 females the second premolar 
was absent, and seen only on one side in 2 animals. By and large dentition, 
especially of the maxillary teeth, is far weaker than in jungle cat — not only 
in length of tooth but also in size. Anterior inner cusp of upper carnassial 
tooth much smaller than in jungle cat and summit far less developed and 
blunter. Canine relatively small but quite powerful and long. 

Skull of female characterized by smaller size, slightly lighter overall 
build, relatively less intense postorbital constriction, relatively close proximity 
305 of semicircular lines (impressions of masseter muscles on roof of skull), 
and characteristics of dentition already described. Age-related changes seen 
in intense swelling of brain case in young animals, relatively short facial 
part of skull, relatively broad interorbital and especially postorbital 
constrictions, absence of crests, and wide distance between temporal lines 
of skull. Geographic variation of skull apparently does not affect its structural 
features, but some insignificant differences in overall size are known. 

Thoracic vertebrae number 13, lumbar 7, sacral 3, and caudal 17-23. 
Length of OS penis in adults 5.0 to 6.0 mm and its width 1.0 to 3.0 mm 
(Haltenorth, 1957). 

Weight of heart in young male (December) 10 g and heart index 4.0%o; 
corresponding values for an aduh male (January) 24.5 g and 436%o. Absolute 
length of intestine in relation to body length in these animals 1,340 cm 
and 1 : 2.43 and 1,500 cm and 1 : 2.42 (Caucasian preserve; based on data 
of Kotov and Ryabov, 1963). 

Diploid number of chromosomes 38. 

Body length of males 430-910 mm and of females 400-770 mm; tail 
length in males 230-405 mm and in females 180-350 mm; length of hind 
foot 120-160 mm and height of ears about 60-70 mm.^^ 

Greatest length of skull in males 96.0-112.3 mm and in females 
87.0-106.6 mm; condylobasal length in males 86.0-102.6 mm and in females 
80-100 mm; zygomatic width in males 62.4-80.0 mm and in females 
60.6-69.0 mm; interorbital width in males 16.5-22.0 mm and in females 
16.0-21.5 mm; postorbital width in males 31.1-36,0 mm and in females 

■'■'Based on 221 animals from various parts of the range of the group, mainly from 
western Europe, including 9 from the Caucasus and 29 from the Ukraine (evidently, western); 
males 137 and females 84 (Suminskii, 1962). Some of the low values evidently pertain to 
juvenile animals. 


30.0-36.5 mm; and length of upper tooth row in males 30.5-33.8 mm and 
in females 26.7-33.8 mm."^ 

Data in literature on weight conflict and are often highly exaggerated. 
The weight of males (185) in Europe reportedly ranges from 3.5 to 15.0 
kg and of females (55) from 3.0 to 10.0 kg (Suminskii, 1962). It has even 
been stated that the weight can exceed these values, reaching 18 kg. The 
source of information about such gigantic animals is the usual hunter's 
exaggerations being published in journals and conscientiously collected by 
compilers. The existence of very large specimens, perhaps heterotic hybrids 
with domestic cat, is not denied, but none attains such sizes. No type of 
food can possibly fatten a cat to a weight of 15 kg. It is significant that 
in these same publications such divergences and exaggerations are absent 
for skull sizes and even body measurements. The normal weight of adult 
males is 5.0 to 6.0 or 7.0 kg; females are somewhat lighter. Cats weighing 
8.0 kg are quite rare in the Caucasus (Dinnik, 1914). In Moldavia cats weigh 
up to 8.0 kg with a body length of up to 830 mm; females weigh up to 
6.0 to 6.5 kg and have a body length of up to 750 mm (Lozan and Korchmar', 
1965). Weight varies perceptibly with age, season of the year, and vagaries 
of environment; when rodents are plentiful, cats are better fed and their 
average weight higher while, given poor conditions, cats can become severely 
emaciated (for more details see "Geographic Variation"). 

Some geographic variation does occur in measurements and weight 
but it is not significant. (V.H.) 

Systematic Position 

See above, under notes on the genus on p. 398, and below in the section 
pertaining to steppe cats (p. 455). 

306 Hybridization of Wild with Domestic Cats, and 

Hybrid Populations 

Reports exist about the hybridization in nature of forest wildcat and domestic 
cat over much of the range of the former. This problem, of significant the- 
oretical interest, is also of practical importance; distinguishing pure and 
hybrid individuals. 

The forest wildcat group can be considered forms which evolved 
autochthonously in Europe during the Early to Middle Pleistocene. The 
ancestral form was F. hmensis Martelli (Villafranchian; Kurten, 1965). The 
domestic cat itself represents the domesticated so-called bay cat of Egypt, 

■''^Animals from the Caucasus (Zoological Museum, Moscow University) with the 
addition of some extreme values taken from European literature (Ognev, 1935; Haltenorth, 
1957; Suminskii, 1962; and others). 


a subspecies of another group of forms of the species (libyca, i.e., F. s. 
libyca). The forest wildcat of Europe (group silvestris) is not in the direct 
line of evolution of domestic cat (see "Origin of Domestic Cats" under 
"B. Group libyca"'). 

The first appearance of domestic cat in Europe was recorded in the 
fourth century A.D. (Rome). For a long time this animal was a rarity, but 
gradually it colonized all Europe (tenth century in England and twelfth 
century in Kiev; Pidoplichko, 1956). There is no doubt that throughout the 
Middle Ages and for much of the later period, in spite of intensive felling 
of forests ("epoch of forest clearing" in western and central Europe in 
the eleventh and twelfth centuries), conditions for the existence of forest 
wildcat were totally favorable everywhere and the population structure was 
normal. For these reasons, and also the small size of the population of 
domestic cat, hybridization with the latter was not significant. 

Rapid deforestation of western and central Europe and afforestation, 
especially in the nineteenth century, coincided with the intensification of 
pursuit and killing of predators (wolf, lynx, and bear) in the interests of 
promoting game. Wildcats were also victims of extermination and their 
range shrank rapidly and their population dropped dramatically. Almost 
throughout the territory of western Europe, apart from some eastern regions 
(Carpathians), these animals had disappeared or reduced to rare (extremely 
rare) animals by the early part of the present century. 

Under these conditions ("lack of partners" in particular), some zoolo- 
gists feel that hybridization with domestic cat, stragglers, or even those which 
had turned feral, became a widespread phenomenon, instead of occasional 
as at the beginning of the prior century. Several zoologists have suggested 
the possibility of spontaneous hybridization. According to another viewpoint 
(Suminskii, 1962 and 1962a) in central and westem Europe hybridization 
was so extensive that, at present, almost all forest wildcats there exhibit 
features of the domestic cat to some extent or the "pure" forest wildcat 
is in general absent in Europe. In any case there are no "pure" populations, 
although the level of admixture of the blood of domestic cat in different 
populations obviously varies — even to the extreme extent of predominance 
of features of domestic cat. Thus the average "percentage of purity "^"^ 
based on craniological features in the French and Swiss Alps is 44.1%, 

^^ According to this author the degree of "purity" is detennined from several features 
(he suggests more than ten), each of which has been assigned a range of values on a point 
scale. Their total in a given skull and skin determines the degree of "purity" of the individual 
with respect to the forest wildcat. The average of these values provides an idea of the ' ' purity" 
of the population. Suminskii 's scheme is far from convincing since the selection of features 
is largely arbitrary, all have been assigned equal value, and the range of their individual 
and age-related changes has not been established (for a detailed critique of Suminskii "s scheme, 
see Sladek, 1966, and especially ICiatochwil and Kj-atochwil, 1970). 


Hungary 61.6%, former territory of Germany 63.1%, Scotland 66.1%, and 
Poland 73.0% (Suminskii, 1962 and 1962a). 

Since the fertility of hybrids is considered low, the destruction of 
wildcats in Europe has been proceeding not only as a result of their pursuit 
and "depriviation of landscapes", but also of hybridization. Thus, in spite 
of measures taken for wildcat protection in some countries, these animals 
are becoming extinct in Europe, although individual "wildcats" have 
continued to survive and could possibly build up the population under proper 
conservation laws. 
307 The above situation prevailed more or less in western Europe (see 

further) but not in the Soviet Union. Conditions for the survival of the animal 
and its population density in the western regions of the Ukraine, particularly 
in the extensive Сафа1Ыап forests, are such that they enabled the population 
to preserve a high degree of purity, although instances of hybridization are 
not excluded. This is even more applicable to the Caucasus. At some places 
there are reports of domestic cats straying or turning feral but offspring of 
wild and domestic cats have mainly been observed in small remote forest 
villages, for example in forest cordons.^"^ However, the regions of 
extensive forests in the Great Caucasus and some parts of the Trans-Caucasus 
are such that hybridization there was practically absent or only a rare 
occurrence. In any case it could not have influenced the character of the 
wild population or its purity.^"^ While evaluating this phenomenon it 
should also be kept in mind that the very possibility of female mating with 
domestic males under conditions of normal density and structure of 
population of wildcats, as prevailing in the Soviet Union, is perhaps close 
to nil. Domestic males undoubtedly cannot successfully compete with wild 
males. Such hybridization can affect the type of domestic cat, which evidently 
did occur (see below), but not the population type of the wildcat. 

In the light of recent data the actual position in western and central 
Europe is not exactly that outlined on the basis of the material studied. It 

^^hus for the Caucasian preserve, V.A. Kotov and L.S. Ryabov (1963) have shown 
"innumerable instances of mating (of wildcat) with domestic cats. We have seen many such 
hybrids in forest cordons .... Such kittens exhibit the body form, coloration, and habits of 
the wild parent. They take fright at the slightest provocation, do not permit fondling, and hide 
the moment strangers appear. As they grow up, such hybrids usually attack domestic birds." 

■'^Suminskii 's statement (1962) that, according to external characters the "percentage 
of purity" ■ of Ukrainian cats (evidently from western regions and the Сафа1Ь1ап8; 29 animals) 
was 51.6% and that of Caucasian (9 animals) 55.7%, does not appear correct in the light 
of the foregoing discussion. The discrepancy is underscored by the fact that this index was 
50.7% for the former territory of Germany, 45.5% for France (16 animals) and 48.7% for 
Poland (18 animals). In these countries hybridization is, of course, far more probable. Such 
contradictions in data underscore either a deficiency in the method itself or inconect selection 
and evaluation of indicator characters. 


appears that in spite of all the unreliable distinguishing features and their 
combinations reported by various authors, forest wildcats and mixed-breed 
domestic cats (wholly domestic, wanderers, ferals, and solitary) do reliably 
differ in volume of brain case, i.e., in size of brain. This characteristic has 
been detected on the basis of the principle established long ago; namely, 
that the brain of domestic cat is invariably smaller than that of its wild 
ancestor (Юatt, 1912). Although the domestic cat has not descended directly 
from the European wildcat, both belong to the same species. This relatioaship 
is maintained in spite of the fact that the cat has undergone less changes 
during domestication than other species. ^^ 

In the forest wildcat the volume of the brain case exceeds 35 cm^, 
while it is less than 32 cm^ in the domestic cat. When the volume varies 
between 32 cm^ and 35 cm^ (rare specimens), these two forms are distin- 
guished by the ratio of the "maximum skull length to the volume of the 
brain case." If this index is less than 2.75 the skull belongs to a wild cat; 
if greater, the skull is that of a domestic cat (index true in 99% of 139 
skulls of wild and 317 domestic cats recovered from France, former German 
territory, Romania, Yugoslavia, Hungary and Switzerland; Schauenberg, 
1969). This critical index for the west Caфathian population could be 
reduced to 2.60, while the lower limit of the skull volume of forest wildcats 
is taken as 36 cm^. In all skulls with a volume index characteristic of 
domestic cats the depression (glabella) at the juncture of the nasals and 
frontals is very prominent. This is a characteristic feature of domestic cats 
(50 skulls of wildcats from western Caфathians and 63 skulls of domestic 
cats which had become feral; central Moravia-Czechoslovakia; ICratochwil 
and Kratochwil, 1970). 
308 Thus it is evident that the European cat populations under study are 

essentially pure. This does not exclude the occasional appearance of indi- 
viduals with some amount of admixture of blood of domestic cats. However, 
it cannot be said that hybrid populations and the "disappearance" of wildcats 
in Europe occurred in this manner. Such an assumption is even less relevant 
to populations of the Soviet Union, although the appearance of some hybrid 
animals, primarily perhaps in Moldavia, cannot be ruled out. However, this 
problem should be resolved in each individual case, apart from obvious 
ones (' 'domestic " ' color, etc.), based on the volume index of the brain case. 

Hybrids of wildcats (F. 5. silvestris and F. s. caucasicd) and steppe cats 
{F. s. caudata) have evidently not been found in nature. The reason perhaps 
lies in the absence of extensive and close geographic and biotopic contact 

•^^Features suggested by various zoologists for differentiating these tw'o forms of cats, 
in addition to other details, apart from articles by Suniinskii cited in the text, can be verified 
in works by Ognev, 1935; Rohrs, 1955; Haltenorth, 1957; and Sladek, 1966. 


between them. In the Soviet Union such hybrids have not been reported 
and apparently do not occur.^^ C^-H.) 

Geographic Distribution 

Found in the forest regions of central and southern Europe, Asia Minor, 
and the Caucasus. 

Geographic Range in the Soviet Union 

The range in the Soviet Union (reconstructed) represents the eastern and 
southeastern segment of the range of the group. It occupies the western 
part of the country and is divided into two sections — European and 
Caucasian (Fig. 184), which are continuous in the south outside the borders 
of the Soviet Union. 

The following were the boundaries of the European part of the range 
in the period of its maximum development over the historic past. In the 
northwest the range covered Lithuania and southern Latvia roughly to the 
latitude of Riga, and northeastern Belorussia to the upper Dnepr in the 
Mogilev district, but apparently a little to the east (former Mogilev gover- 
nance). Farther on the boundary turned southeast, encompassing part of 
former Kaluga governance, former Voronezh governance, and in particular 
Shipov forest on the Don at Pavlovsk. From there the boundary evidently 
proceeded southwest toward the great bend of the Dnepr, extending south 
of Zaporozh', and descended to the sea along the Dnepr. To the west of 
this line the range everywhere extended to the western Soviet borders and 
into central and southeastern Europe. 

For the region between the above line and the Dnepr, information is 
available about the occurrence of ihis group at Kursk and Belgorod (Kursk- 
Belgorod territory, land of Belgorod polko), Putivl', Sumy Akhtyrka 
(southern Khar'kov), and along both banks of the Dnepr in the land of 
Zaporozh 'e Sechi. Information for these sites pertains to the seventeenth 
and eighteenth centuries and only to part of the nineteenth. It is quite possible 
that some references may have been based on encounters with feral cats,'^^ 
but some definitely pertain to wildcats. It should be kept in mind that 

■^^Haltenorth (1953), citing Ognev (1930), who cited Smirnov (1922) affirms instances 
of hybridization of wild and steppe cats in the Trans-Caucasus. This is erroneous; both Ognev 
and Smirnov mention only hybrids of domestic and steppe cats. 

''•'Such, for example, was the report pertaining to the "former Usmansk county of 
Voronezh governance (Kjiyazhinskii, 1924, according to Temovskii, 1929) in the present 
century. Along with this, the eyewitness account of A.P. Volynskii who wrote in 1736 and 
1738 that wildcats of "a genus distinct from Astrakhan cats" existed in Voronezh governance 
(Kirikov, 1959) is wholly trustworthy. 


conditions for the survival of wildcats in the territories under consideration, 
some 200 to 300 years ago, were far more favorable than at present. This 
is particularly true of the region between the Dnepr and Northern Donets 
and the Don basin. This was particularly so during the remote past (Middle 
Ages; Heptner, 1959). In any case the outline of the range in the Russian 
plains is quite typical of forms of western origin. 

References by some older zoologists to the occurrence (cited from 
Ognev, 1935) of cats in the former Belozersk county of Novgorod gover- 
nance, former Vytegorsk (Vishegorsk in Ognev, 1935 is a misprint) county 
of Olonets governance, former Shenkursk county of Arkhangel'sk 
governance, and finally to the southern (Zlatoustovsk) Urals are neither 

309 tmstworthy nor confirmed. Intrusions at medieval Pskov (eleventh to fifteenth 
centuries; Tsalkin, 1952) pertain not to wild but obviously to domestic 

In the last century the range shrank to insignificant remnants. This occur- 
red probably due to changes in environmental conditioas and the general 

310 scarcity of game on which wildcats feed. It was not exposed to special direct 
pursuit. The chronology of this process is not clearly understood. Thus it 
has been suggested that all the data on wildcats of Latvia and probably 
also of Lithuania for the nineteenth century and possibly even the end of 
the eighteenth, pertain not to wildcats proper but to domestic cats turned 
feral (Paaver, 1965). In any event, cats disappeared earlier in Latvia than in 

In the stage of range contraction preceding the present one, i.e., in the 
nineteenth century, at places even in the present century, the eastern boundary 
commenced evidently somewhere in Belorussia, in the Minsk or Mogilev 
region. To the south the easternmost points of habitation of forest cat extend- 
ed into the Kiev district to the west and southwest of Юev (Radomyshl', 
Stavishchi to the south of Belaya Tserkva), although it was evidently 
encountered right in Kiev. From these places the boundary turned in the 
direction of the Podol'sk district where the forest cat was recorded 
particularly in the region of Ushitsa in the basin of the upper Dnestr. From 
there the range boundary descended southward, encompassing the region 
between the Dnestr and Danube, to its eastern boundary at the swampy 
estuary of the Dnestr. It is possible that forest wildcats in the nineteenth 
century extended even farther east and lived, for example, in Chemyi forest 
near Юrovograd and in swamps and other places in the Dnepr valley, but 
sufficiently reliable data are not available. 

In the 1940's to the 1960's cats were reported from the western half 
of, and northern (Poles 'e) Volyn district, the extreme north of L'vov, and 
from the northern part (Poles 'e) of Roven and Zhitomir, and probably was 
encountered in the southern part of Zhitomir district and some places in 


20 25 30 35 40 45 50 55 60 65 

•♦•t«».t»t В 

45 Araks R. 50 

Fig. 184. Range (reconstructed) of European forest wildcats, Felis (Felis) sihestris 

silvestris and F. (F.) s. caucasicus (Caucasus) in the Soviet Union. 
A — boundary of maximal reconstructed range (sixteenth to the nineteenth 
centuries); В — range boundary in the west in the 1940's to the 1960's (see Fig. 
185 for details of the range in Moldavia). Question mark in the west pertains to 
possible occurrence in the middle of the twentieth century in Izmail, Zhitomir, 
and Kamenets-PodoF districts and in Dnepr swamps. The following are some 
individual places of occurrence in the past: 1 — "bCaluga governance"; 
2 — "Voronezh governance"; 3 — Kursk; 4 — PutivF; 5 — Sumy; 6 — Aktyrka; 
7 — Belgorod (together with No. 3, "Kur-Belogorod territory"); 8 — Shipov forest 
at Pavlovsk on the Don; 9 — Zmiev; 10 — Chemyi forest; 11 — Dnepr .swamps; 
12 — Mogilev. Crosses denote some erroneous references (V.G. Heptner). 


Fig. 185. Distribution of European forest wildcat, Felis {Felts) s. silvestris in 


1 — Kagul swamp (lower Prut); 2 — broad-leaved forest of Kodr, 3 — northern 

Moldavian forest; 4 — floodplain forests and swamps of lower Dnestr (Yu.V. Averin 

and M.N. Lozan — data for 1967). 

Kamenets-Podol'sk. Wildcats were generally found in all the montane forests 
of the Caфathians within the Trans -Caфathian, Stanislav, Drogobych, and 
Chemovits districts, in the greater (southern) part of L'vov, and in the 
southeastern part of Ternopol 'sk (Tatarinov, 1956). 

In Moldavia forest wildcats are distributed in forests along the Prut 
(almost to the mouth), at places along the Dnestr, and interfluve forests 
in the so-called Kodr (in Strashensk, Orgeevsk, Teleneshtsk, and Kotovsk 
regions; Kuznetsov, 1952; Yu.V. Averin and M.N. Lozan; Fig. 185). 
Evidently they also live in Dnestr swamps. There is no information relating 
to the far south of the interfluve of Dnestr and Danube (Izmail' district); 
occurrence is generally shown, however, as "swamp forests" of the Dnestr 
and Danube (Sokur, 1960).4i 

■^^The map given by this author does not confirm these areas, referred to in the text 
as occurrences in the Dnepr estuary. 


Forest wildcats have not been reported from Belorussia during the 
present century. Only one intrusion has been recorded for Minsk, in 1927. 
The animals are not even seen in Belovezh Forest where they were very 
common at some places. However, their appearance is still possible in 
southern Poles 'e from adjoining sections of Volyn, Roven, and Zhitomir 
311 In the Caucasus wildcats are distributed extensively in all forest regions 

of the Great and Little Caucasus. In the latter they occur in the east to Shusha 
meridian; they are absent only in the extremely high mountains, in the 
forestless mountains of Dagestan, and in the open steppes and semisteppes 
of the eastern Trans-Caucasm. In the Cis-Caucasian plains forest cats are 
present along the forested sections of the Kuban valley and encountered among 
reeds of the Kuban delta. They likewise live in forests (Paraboch) and shrubby 
regions along the Terek, at least to Kizlyar, but probably to the sea. 

The animal has been sighted in the Kuma valley down to its middle 
course, in the Sulak valley, and in the forests of Stavropol'. 

A reference to the presence of the forest cat in Talysh (Radde, 1886) 
has been questioned (Satunin, 1905; also Smimov, 1922) since the skull 
brought from there proved to be that of domestic cat. Yet even now (Bobrin- 
skii et al., 1944 and 1965; and others) wildcats are nonetheless reported 
for the Talysh and Lenkoran. However, since the 1880's no sightings other 
than the first report mentioned above are known from this part of the 
Caucasus and the animal was/is evidently absent there. Only the jungle 
cat is definitely known to occur in this region. 

The range of the forest wildcat in the Caucasus underwent no significant 
changes in the period under consideration. Thus, in spite of a reduction 
in forests in the Stavropol' upland, cats still lived in Temnolessk forest 
40 km south of Stavropol'. Only localized reductions occurred due to 
deforestation. However, this species is also found in gardens and vineyards 
with sparse forests and shrubby thickets (Kizlyar for example).'^^ 

Geographic Range outside the Soviet Uniofi 

The range outside the Soviet Union (see Fig. 181) encompasses Europe in 
the west, including England (absent in Ireland and evidently never present 

''-Range according to Bikhner, 1905; Satunin, 1905 and 1915; Dinnik, 1910; Brauner, 
1914, 1923, and 1928; Migulin, 1928; Charlemagne, 1928; Temovskii, 1929; Fedyushin, 
1930; Ognev, 1935; Heptner and Formozov, 1941; Vereshchagin, 1947 and 1958; Kalnin'sh, 
1950; Heptner, Turova and Tsalkin, 1950; Kirikov, 1952, 1959, and 1966; Komeev, 1952; 
Kuznetsov, 1952; Serzhanin, 1955 and 1961; Tatarinov, 1956; Sokur, 1960 and 1961; Lozan 
and Korchmar', 1965; Paaver, 1965; Alekpeiov, 1966; Korchmar', 1968; and also original 
data of Yu.V. Averin, M.N. Lozan, V.G. Heptner, and others. 


there), northward to the Baltic Sea, and southward to southern Spain, Italy, 
Sicily, the Balkans, and all of Asia Minor. These cats are not known to 
occur in Syria and Iraq but may perhaps intrude into areas adjacent to Turkey. 
They have not been reported for Iran and are evidently absent there. Cats 
inhabiting islands of the Mediterranean Sea (Corsica, Sardinia, Crete, and 
the Balearic Islands), sometimes even those of Italy and Sicily, are placed 
in this group, and/or libyca group, or considered a transitional group. 

Finds of Pleistocene cats assigned to this species generally do not occur 
beyond the limits of the present-day range. Wildcats have been found in 
Holocene Crimea (Birulya, 1930)* and in cultural horizons of some five 
thousand years ago in southern Scandinavia (Scania; Eckman, 1922)", and 
probably extended to the Gulf of Finland in the Pri-Baltic during the 
Holocene (Paaver, 1965)*. The depiction of the Holocene ("post-glacial") 
range as having encompassed the eastern part of the Don basin and central 
Volga, including also Trans- Volga (Haltenorth, 1953) is evidently based 
on assumptions. There is no concrete proof of this range extension in our 

In all probability forest wildcats were not distributed beyond the Don 
bai^n and perhaps not even its western part. In any case cat remains from 
Sarkel (lower Don, ninth to the twelfth centuries) have been adjudged 
those of domestic animals (Vereshchagin, 1959). Evidently European and 
Caucasian portions of the range may be regarded as not contiguous across 
the Russian plains. This is indicated by the independent status of the 
subspecies of the Caucasian form and its affinity to animals from Asia Minor. 
Forest wildcats intruded into the Caucasus from Europe, where they were 
autochthonous (Kurten, 1965), through the Balkans and Asia Minor in the 
Pleistocene. (V.H.) 

312 Geographic Variation 

Several subspecies have been described for the forest wildcats of the group 
defined here; however, they have not been satisfactorily characterized to 
date, and have been separated for the part without adequate basis. Although 
geographic variation within the Soviet Union still requires clarification, two 
forms may be tentatively recognized in the USSR. 

1. Central European forest wildcat, F. (F.) s. silvestris Schreb., 1777 
(syn. ferus, ferox, and euxina). 

General color quite dark with a gray tone; pattern on head, dorsal spinal 
band, and transverse stripes and spots on trunk distinct and usually vivid. 
Rings on tail may range from three to ten, but more often seven in number. 

*Not in Literature Cited — Sci. Ed. 


Body length (20 animals; western Ukraine, Caфathians) of males 
480-910 mm (M 633) and of females 400-690 mm (M 540); tail length 
in males 330-350 mm (M 313) and in females 280-340 mm (M 294) 
(Suminskii, 1962y^ 

Data on the measurements of forest wildcats from Moldavia and adjoin- 
ing places are scanty and somewhat contradictory. The average body length 
of three males from Moldavia has been reported as 810 mm and of four 
females as 750 mm; tail length in males 360 mm and in females 355 mm; 
length of hind foot in males 160 mm and in females 124 mm; height of 
ears in males 60 mm and in females 49 mm; and weight of males up to 
8.0 kg and of females to 6.5 kg. 

Greatest length of skull in males (three) 87.8-91.8 mm; condylobasal 
length of skull in males (five) 81.2-104 mm and in females* 80-91 mm; 
and zygomatic width in males (six) 62-78 mm and in females* 61-90 mm 
(Yu.V. Averin and M.N. Lozan; Mindlin, 1938)**. 

According to other data (Brauner, 1928) animals from former Bes- 
sarabsk govemance, former Odessa county (one specimen from Podolia) 
had the following measurements (males and females together): body length 
(eight) 520-630 mm (M 590); tail length (eight) 310-340 mm (M 320); 
length of hind foot (five) 125-140 mm (M 133); and height of ears (five) 
60-63 mm (M 61). 

Condylobasal length of skull (six) 83-1 13 mm (M 98.7); muzzle width 
above canines 22-31 mm (M 26.6); interorbital width 31.0-43.2 mm (M 
36.4); zygomatic width 64.0-86.5 mm (M 75.1); and length of upper tooth 
row 21-30 mm (M 26). 

Greatest length of skull (mean, Poland, 10 males and females) 97.4 
mm; condylobasal length 88.3 mm; zygomatic width 69.1 mm; interorbital 
width 19.8 mm; and postorbital width 34.9 mm (Suminskii, 1962). 

Greatest length of skull of fully adult animals from central Europe 
(Alsace and western regions of former German territory): males and females 
(25) 88-110 mm (M 94.1); greatest length of skull in males (6) 96-110 
mm (M 102); condylobasal length in males (26) 86-101 mm (M 91.6) and 
in females* 80-S9 mm (M 84.7); and zygomatic width in males (26) 66-80 
mm (M 71.9) and females* 63-69 mm (M 65.3). 

Weight: males and females (189) 3,750 to 11,500 g (M 6,880); males 
(108) 5,000 to 11,500 g (M 7,180) and females (33) 3,750 to 10,000 g 

''-■'Information on this cat is almost nonexistent in Soviet museums and data in Soviet 
literature scanty. Information derived from foreign literature and from adjoining or nearby 
territories has been used to characterize it. Data by Suminskii (1962) on weight are extremely 
exaggerated (males 5.9 to 14.8 kg, etc.) and have not been included here. 

*Number of females omitted in Russian original — General Editor. 

**Not in Literature Cited — Sci. Ed. 


(М 5,970) (Haltenorth, 1957). Although these figures do not include extreme 
values (see above), they are nevertheless sometimes considered, not without 
reason, to be on the high side. High values are usually attributed to hunters' 
errors. Thus the weight of 36 pure-blooded adults, in part extremely large 
animals from Slovakia, weighed and measured by a zoologist (Sladek, 1966) 
was: males (20) 3,300 to 7,700 g (M 5,248) and females (16) 2,600 to 5,840 
g (M 4,233). The largest male in this series had a body length of 880 mm, 
tail length 310 mm, length of hind foot 145 mm, and height at shoulders 
430 mm. Greatest length of skull was 107 mm, condylobasal length % 
mm, and zygomatic width 76 mm. 

Found in the extreme west of the European part of the Soviet Union. 
313 Outside the Soviet Union this form is encountered in the European 
part of the range of the species, except in Scotland and islands of the Mediter- 
ranean Sea. 

2. Caucasian forest wildcat, F. (F.) s. caucasica Satunin, 1905 (syn. 
daemon and trapezia). 

General shade of color gray, on the average somewhat lighter than in 
nominal form; pattern on head and dorsal band well developed, but pattern 
on trunk and partly on tail reduced. Transverse bands and spots on trunk 
usually faint or absent, but well defined in a few individuals. In addition 
to black tip on tail, only three distinct black transverse rings usually present. 

"Pelage rich, dense, and soft. On the back an average of 8,900 hairs 
occur in a 1 cm^ area, and 2,000 in the same on the abdomen; ratio of 
underfur hair to top hair, 10:1. Guard hair stand somewhat above top hair. 
Latter comparatively short and do not rise much above underfur. Average 
length of guard and four categories of top hair on the back: 44, 41, 39, 
37, and 36 mm, and thickness 81, 87, 71, 62, 43, and 20 microns; underfur 
hair 34 mm and 20 microns respectively. Hair length on abdomen less than 
on back" (B.F. Tserevitinov).^^ 

Body length of the adult cats usually 70 to 75 cm, height at shoulders 
26 to 28 cm, and height at sacrum 30 to 32 cm. Weight often 5,200 to 6,000 
g and up to 8.0 kg; rarely even more (Dinnik, 1914; Satunin, 1915). Two 
adult males from Caucasian preserve weighed 5,610 to 6,200 g and three 
females between 4,150 and 5,000 g. 

Found in the Caucasus. 

Outside the Soviet Union, found in Asia Minor. 

Melanistic individuals are encountered among Caucasian cats and have 
been separated into a distinct species, F. daemon. Actually these are simply 

''''The information given here differs significantly from data given above for central 
European cats. This discrepancy is evidently due more to differences in methodology than 
to differences between the two subspecies. The data of B.F. Tserevitinov in any case are 
wholly comparable to that given for other Soviet cats. 


feral domestic cats (material from the Zoological Museum, Moscow Uni- 
versity) or, perhaps, hybrids of wild and domestic cats. Occasional cases 
could well be melanistic wildcats, since some geographic localization of 
these forms is seen in the eastern part of the southern slope of the Great 
Caucasus in the districts of Lagodekh, Zakatal, and Nukhi (former Nukhinsk 
314 county and former Zakatal 'sk area). Data are available for the last century 
about the presence of such forms in the eastern part of the Little Caucasus 
from Kirovabad (former Elisavetpol') to the upper Terter and Shusha 
(Satunin, 1915). 

In the middle of the present century they have generally been caught 
in the same places in Little Caucasus, namely, Stepanakertsk and Shushinsk 
regions of Azerbaidzhan (Karabakh range, two to four animals in winter; 
Alekperov, 1966). In 1948 in a fur warehouse in Baku, of the 117 skins 
of forest wildcats 14 were pure melanistic and 3 dark brown (A.P. Komeev). 
Evidently these figures represent all of Azerbaidzhan. 

The Caucasian wildcat is a distinct well-characterized form and its range 
readily demarcated from that of western forest forms. Unfortunately a 
comprehensive taxonomic analysis has not been attempted to date. However, 
it has always been thought that this form is larger than the European form, 
which is not correct. The main difference between the two lies in the color 
characteristics described above. 

One of the features of the Caucasian form, compared with the western, 
is that no hybridization of the former with domestic cat takes place, which 
is a very common occurrence in the western form. There is no information 
from the Caucasus about extensive hybridization, let alone hybrid popu- 
lations. Living conditions are not favorable for hybridization. Extensive 
development of suitable landscapes and extensive forests (Kuban Caucasus, 
and a series of other areas) ensure a natural population in numbers and 
composition. Figures relating to the admixture of blood of domestic cat 
in the Caucasus (see above) are no doubt erroneous and reflect imperfect 

Outside the Soviet Union, apart from the two forms described above, the 
following subspecies of European forest wildcats are usually recognized: 
1) F. (F.) s. grampia Miller, 1907— Scotland; 2) F. (F.) s. tartessia Miller, 
1907— southern Spain; 3) F. (F.) s. molisana Altob., 1921— Italy; 4) F. 
(F.) s. euxina Pocock, 1943 — Dobruja; and 5) F. (F.) s. morea Trouess., 
1904 — southern Greece (Ellerman and Morrison-Scott, 1951). Only the 
subspecies grampia would appear genuine (Haltenorth, 1957). 

Thus there are only three subspecies of forest wildcats. The view that 















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there can be no geographic variation due to hybridization of forest wildcat 
with domestic cat is not correct (Suminskii, 1962). 

The forms listed above are typical for the group silvestris. The following 
forms occupy an intermediate (transitional) position in relation to the libyca 
group: 4) F. (F.) s. jordansi Schwarz, 1930 — Balearic Islands; 5) F. (F.) 
s. reyi Lavaud, 1929 — Corsica; 6)F. (F.) s. cretensis Halt., 1953 — Crete; 
and 7) F. (F.) s. sarda Lat., 1885 — Sardinia, Sicily, northern parts of the 
North Africa from Cyrenaica to Morocco and southern Atlas, and Algerian 
Sahara (Haltenorth, 1953 and 1957). 

Often, all of these forms are combined into one (sa?-da) and placed 
in the libyca group (species). (V.H.) 


Population. In the western parts of the range, the forest wildcat became 
scarce even in the early nineteenth century in Latvia and almost extinct 
in Lithuania and much of Belorussia. They are, at present, extremely rare 
and only single individuals are caught in Lithuania and Belorassia (Poles 'e). 
About 1,000 cats still live in the Carpathians and over a hundred are 

Fig. 186. Montane forest in the Carpatiiians. Habitat of forest wildcat and lynx. 
Vicinity of town of Rakhovo. August, 1967. Photograph by G.I. Simkin. 



caught annually in the Trans-Carpathians and the western part of Volyna; 
cats are now extinct elsewhere in the Ukraine. In Moldavia forest wildcats 
are common in the lower reaches of the Dnestr and Prut but rare in the 
Kodr. In the early 1960's from 20 to 40 cats used to be caught every season 
in this region. 

Forest wildcats are quite common over large areas in the Trans-Caucasus 
and the Caucasus, in the north to deltas of the Kuban and Terek. They are 
particularly numerous along the Black Sea coast but rare near the Caspian 

315 Fig. 187. Ancient beech forest. Habitat of forest wildcat in Babich area. Carpathians. 

Photograph by Yu.D. Tret'yak. 


[Sea]. In Azerbaidzhan these animals are caught in the thousands, in 
Armenia, Georgia, and Dagestan in the hundreds, and in the Groznen district 
and Krasnodar territory in the northem Caucasus again in the thousands 
(up to 5,000). In the forested "Chemye Gory" range — the first ridge of 
the mountains — wildcats were numerous in the central part of the northem 
slope of the Great Caucasus in the 1920's (V.G. Heptner). 

Habitat. In the westem regions of the Ukraine wildcats inhabit broad- 
leaved hornbeam-oak, beech, and mixed forests on the Volyno-Podol'sk 
plateau and in the Caфathians. They are absent in the spruce taiga along 
the crest dividing the Caфathians, and the subalpine belt of these mountains. 
Along particular slopes of the mountains they ascend to the upper boundary 
of beech and mixed forests at a height of 1,200 to 1,400 m above sea level. 
The main habitats are beech forests on the southwestern slopes of the 
Caфathians, i.e., the old "preserved" forests growing at the upper boundary 
of wooded vegetation. These forests consist of massive, centuries-old beech 
and sycamore trees, which yield an abundant crop of seeds every two or 
three years. Many trees have hollows in which not only birds but also 
squirrels, martens, and cats nest. In these forests mouse-like rodents and 
birds are abundant; wildcats mainly hunt the first of these (F.I. Strautman; 
Konyukhovich, 1953; Taratinov, 1956a). 
317 In Moldavia wildcats inhabit beech forests as well as swamps consisting 
of dense thickets of reeds and tala willow with some hollow willows and 
black poplars growing among them. Such swamps abound in small tributaries 
and lakes. In the swamps of the lower Prut there are no forests in the 
floodplains but wildcats are found there in large numbers. They live in the 
hollows of old willows along dams and on ridges, in reed thickets along 
banks of reservoirs, and on floating islands (Lozan and Korchmar', 1965). 
The habitats of these cats are particularly diverse in the Caucasus. They 
are commonly found on ridges, islands, and in dense reeds in the swamps 
of the Kuban and Terek, and quite often encountered on islands that have 
broken off and are floating far from shore. There it leads a life similar to 
that of jungle cat. In Dagestan forest wildcats are encountered in reeds along 
the sea coast (Dinnik, 1914) and in Georgia regularly visit marine surf zone 
(M.P. Pavlov). Along the southern slope of the Great Caucasian range, they 
live down to the very shore of the Black Sea where they are confined to 
thickets of cherry laurel, yews, and box trees (A.A. Nasimovich). Wildcats 
are common, at places even abundant, along montane slopes overgrown 
with beech and oak. In the Caucasus they prefer the broad-leaved forest 
zone, especially sections which adjoin the glades and valleys of rivers, and 
inhabit mainly the central montane belt. The cat is met with significantly 
more rarely in dark coniferous forests, and in the subalpine belt it occurs 
only as an exceptionally rare occurrence. Thus in Caucasian preserve 76.7% 


316 Fig. 188. Old beech forest with vines. Habitat of forest wildcat on Black Sea coast 

of the Caucasus. Photograph by A.P. Zhandarmov. 

of occurrences took place in the belt of broad-leaved forests, 13.6% in dark 
coniferous forests, and 9.7% in the subalpine belt. Instances of occupation 
of the deep montane regions covered with coniferous forests are few; the 
cat is confined exclusively to river valleys usually that support broad-leaved 
species (Teplov, 1938). The above distributional features of cats according 
to habitat depend on availability of food, primarily its accessibility during 
the winter period. 

In montane forests cats show preference for cliffe or rocky sections. 
They invariably prefer rock cliffs to hollows of trees and set up their dens 


far more rarely in the latter than among rocks. In the forest, except for rocky 
sections, they select impenetrable thickets and places carpeted with fallen 
318 trees and branches. In shrubs and sparse forests they usually select places 
with outcrops of rock slides, stones, and precipices (Dinnik, 1914). In years 
of mass breeding of rodents in the steppes, cats sometimes descend the 
foothills to catch voles in fields (see below). At such places, inhabited also 
by jungle cat, wildcats live in the "upper" forests along the slopes and 
not in the valleys (Zakatalo-Nukhinsk valley; N.N. Rukovskii). Most of 
the population lives in the mountains at a height of 300 to 600 m above 
sea level and even higher (600 to 800 m) at some places; when intmding 
into the subalpine beh, however, they sometimes ascend up to 1,500 to 
1,800 m above sea level (Dinnik, 1914). In Armenia wildcats live in the 
forest regions at a height of 700 to 2,500 m above sea level (Dal', 1954). 
In some parts of the Caucasus this cat lives in extremely wet places, 
apart from river mouths (see above). For example, in the Kolkhid lowland 
(Batumi and Poti) rains are very frequent, with downpours almost every 
day in autumn and winter. Local cats are not afraid of dampness and wander 
day after day in mud, wet grass, or weeds, which drip water on them. They 
also enter and swim through rivulets and brooks. During high floods, when 
tens of kilometers of lowland forests and swamps are inundated, they live 

317 Fig. 189. Beech forest on the southern slope of Great Caucasus range. Biotope 

of forest wildcat. Azerbaidzhan. November, 1957. Photograph by N.N. Rukovskii. 


in trees for weeks, feeding on Norway rats taking refuge there (N.K. 

A wildcat will not avoid the proximity of humans and often lives not 
only close to settlements but even in granaries, baras, attics, summer houses, 
and larger residences. 

The European forest wildcat is not adapted to living in regions with 
deep, loose snow cover. With a high weight load on the resting surface 
of the foot at 88 to 118 g per cm^, it flounders and advances with great 
difficulty on loose snow since it is relatively short-legged (height of forelegs 
and lower surface of chest to the ground in males 25 cm, and in females 
22 cm). Moreover, it cannot get at rodents under the snow. These are 
evidently the reasons why the cat does not ascend high into the mountains, 
especially in winter. In Caucasian preserve with a deep snow cover the 
cat descends the mountain slopes, avoiding regions of deeper, more abundant 
snow depth which are difficult to negotiate and where birds are scarce 
319 (A.A. Nasimovich). The wildcat usually inhabits only regions where the 
mean of winter snow cover does not exceed 10 to 20 cm (Formozov, 1946). 
This could explain its absence in the central zone of European Russia (depth 
of snow cover 40 to 60 cm), not to mention more northerly regions. 

318 Fig. 190. Thickets of sea buckthorn, with common buckthorn, blackthorn, and 

dog rose. Habitat of forest wildcat. "Chernye Gory" south of Ordzhonikidze, 
near Balta. May, 1966. Photograph by I. P. Mitina. 


In unusually severe, abundantly snowy winters forest wildcats suffer 
starvation and sometimes even die of emaciation. For example, in the winter 
of 1953/1954, in the Kuban and in all of the southern European part of the 
USSR, there was unusually severe, prolonged and constant subzero 
temperatures together with excessive snowfall. The minimum air temperature 
was -21° С in November, -24° С in December, -26° С in January, and 
-35° С in February. Snow drifts in swamps were 2.0 m deep and at places 
even more. This winter proved fatal to many inhabitants of these swamps. In 
Febniary, exhausted and emaciated wildcats, foxes, and wild boar were 
observed to abandon their usual caution and moved onto farms during the day; 
cats often attempted to hide in cattle sheds. Not only a large number of small 
animals and birds died, but predators also suffered from hunger and died 
of emaciation. In March, 1954 one wildcat, three foxes, and two minks 
[Mustela lutreola] were found dead (Pavlov and Kiris, 1956). Wildcats often 
endure starvation and die of emaciation during winter even in Central Europe 
(Haltenorth, 1957). In the very severe winter of 1962/1963 a deep snow cover 
deprived wildcats, especially those living in montane regions, of mobility, 
crippling their search for rodents. They descended into farmsteads. In February, 
1963, dead cats with empty, stomachs were found (Miiller-Using, 1963). 
Domestic cats are capable of surviving total starvation for 19 days. 

Food. In the Carpathians wildcats mainly feed on the yellow-throated 
mouse [Apodemus flavicolis], red-backed vole [Clethrionomys rutilus] and 
ground vole [Pitymus subterraneus] which occur in large numbers during 
"mouse" years in beech forests (F.I. Strautman); they also catch field hares 
[L. eiiropaeus] (Tatarinov, 1956). In the Tгans-Caфathians they feed mainly 
on mouse-like rodents, galliform birds, and squirrels (Konyukhovich, 1953). 
In the Dnestr swamps they hunt for small voles [Mic?otus], water voles 
[Arvicola], and birds (Brauner, 1923); in the Prut swamps, judging from 
320 an analysis of 77 food samples, they feed mainly on water voles (91% of 
samples, Norway rats (24%), and muskrats (13%). Of the birds available, 
they most often attack warblers (52%), white-eyed pochards [Aythya nyroca] 
(20.8%), coots [Fulica atra] (9.1%), spotted crakes [Porzana porzana] 
(5.2%), and gadwall ducks [Anas strepera] (2.6%). Moreover, they eat wood 
mice [Apodemus sylvaticus] (3.9%), fish (3.0%), moliusks (2.6%), and 
beetles (1.5%). In Moldavia, in the lower courses of the rivers, rodents 
predominate in the winter diet while birds, fish, and crayfish predominate 
in the summer diet. Wildcats catch a very large number of aquatic and other 
birds in May and June when most of these birds are sitting on their nests 
(Lozan and Korchmar', 1965). 

In the Kuban delta the main food of wildcats consists of Norway rats 
and water voles, which are abundant there, and additionally muskrats and 
waterfowl (M.P. Pavlov and L.A. Volyanskii). In the northern Caucasus the 


319 Fig. 191. Thickets of willow, blackbeny, blacklhom, and reeds. Biotope of forest wild- 

cat in Terek valley, northwest of Groznyi. May, 1966. Photograph by I.P. Mitina. 

main prey of this cat in summer are mouse-like rodents and edible dormice 
[Glis glis], and more rarely birds. From time to time various fruits, including 
mountain ash, have been found in stomachs. On rare occasions it will attack 
young roe deer and chamois (Dinnik, 1914). In Dagestan broods of pheasants 
and partridges suffer most from this predator (Heptner and Formozov, 1941). 
On the Black Sea coast, judging from an analysis of food remains, it feeds 
on small birds, mice, shrews, and hares. On one occasion the feathers of 
a white-tailed eagle [Haliaetus albicilla], and on another the skull of a kid, 
were found in dens. When necessary, it eats a fresh kill but usually 
approaches it only after jackals have begun to devour it. With the appearance 
of a wildcat, jackals withdraw from the carcass and retum to it only after 
the cat has left (Obraztsov, 1928). 

In Caucasian preserve (12 stomach contents and 35 feces, as percentage 
of total number of both) the food of wildcats is as follows: mammals 100, 
rodents 95.7, mouse-like rodents 93.6, bush voles [Pitymys] 76.6, wood 
mouse 14.9, edible dormouse 2.1, European hare 2.1, mole [Talpa] 4.2, shrew 
4.2, chamois 2.1, small birds 3.5, insects 2.1, and sedge and grass (leaves) 


10.6 (Teplov, 1936). Chamois remains were found in feces collected on 
December 22; the cat evidently ate carrion. Similarly, remains of a drake 
mallard eaten by a cat were also seen. The stomachs of three cats caught 
in August and September in Zakatalo-Nukhinsk valley contained yellow- 
throated mice and edible dormice; the remains of a rock partridge were 
found near a den. In the stomachs of cats caught in summer in the Ismaillinsk 
region of Azerbaidzhan only the remains of yellow-throated mice, wood 
mice, and edible dormice were found (Rukovskii, 1953). 

During the drought of 1948 many wildcats and jackals were sighted 
on the Black Sea coast around drying lakes (for example, Lake Imanati, 
upper Gurinki River), which supported a large number of nutrias and aquatic 
birds. These predators regularly hunted for rodents and birds. In January, 
1949, after the freezing of pools unaffected by drought, the cats turned to 
nutrias hiding in bushes in large numbers as they were compelled to live 
on land. In 1939 in Abkhaz'ya some 60 wildcats and 70 jackals were 
extirpated during the summer over a comparatively small area of the Inkit 
swamp. It is not surprising that with such a huge population of predators 
in Georgia after the severe winters of 1948/1949 and 1949/1950 nutrias 
everywhere became rare. In the stomachs of four cats caught in December, 
1949, near Lake Inkit there were remains of mallards, sheldrakes [Tadorna 
tadorna], song birds, water voles, and small mouselike rodents (M.P. Pavlov). 
Cats caught on June 12, 1937, in Armenia in a beech forest had filled their 
stomach with lizards (E.P. Spangenberg). 

In western Europe the cats feed on rodents (hamsters, Norway rats, 
dormice, water voles, voles, and wood mice) and from time to time on small 
predators such as martens, polecats, ermines, and weasels in addition to 
shrews and moles. They do not relish insectivores. The remains of squirrels 
and forest birds constituted an insignificant quantity in stomach analyses 
(Satunin, 1915; Beaufort, 1961; Traubot, 1961). Wildcats there also attack, 
from time to time, the fawns of [red] deer, roe deer, and chamois. They 
hunt hares and rabbits and catch fish, inflicting some damage during 
spawning, especially on trout (Gaake, 1901*; Dinnik, 1914; Haltenorth, 1957; 
Beaufort, 1961). Individual cats specializing in catching the young of 
321 ungulates are also encountered. When living close to human settlements, 
often right inside them, this predator can cause significant damage to 

From the foregoing data on the food of the forest wildcat in different 
parts of its range it may be concluded that small rodents (mice, voles, and 
rats) constitute their primary food everywhere; birds (chickeas, ducks, and 
more rarely passerines) occupy second place, followed by dormice, hare, 

*Not in Literature Cited — Sci. Ed. 


nutria, muskrat, and insectivores. The predator hunts other animals in years 
of low population of mouse-like rodents, or incidentally. 

While hunting, the cat wanders throughout the forest and along the edges 
of forests and glades. Where conditions are favorable, the forest wildcat will 
readily feed in a field. In the winter of 1923-1924 the weather was free of 
snow in December and January around Vladikavdaz. It was a year of high 
numbers of small rodents, mainly gray voles [Microtus]. Cats, also abundant 
that year, left the forests in the foothills (Chemye Gory), together with foxes, 
during twilight hours and entered singly into fields adjoining the forest. Even 
wolves came. A group of hunters shot several wandering cats and foxes in 
the brief twilight period. The cats were very obese (V.G. Heptner). 

In pursuing prey, sometimes the cat climbs to the very top of high trees, 
even jumping from one tree to another. It lies in wait for a prey and catches 
it by executing a few leaps. Sometimes it lies in wait standing (lying) near 
a burrow or crevice among rocks. Its leaps can reach a span of 3.0 m. The 
cat relies on vision and audition in searching for prey. Olfaction is relatively 
weakly developed. It hunts for nutria and waterfowl from trees overhanging 
the water in which they swim. These cats have been observed hunting in 
Georgia during the dawn hours (Pavlov, 1953). On one drying lake a cat 
attempted to attack an adult nutria feeding in a strongly shoaling and boggy 
section of the shore. The cat approached the nutria by executing large bounds 
but soon mired in the mud and had to return to its former place of ambush 
(M.P. Pavlov). If the prey is small, the cat grabs it in its claws and then 
kills it by piercing the neck or occiput. There are references to this cat 
jumping onto the back of large animals and attempting to bite the neck 
or carotid (Bikhner, 1905; HaUenorth, 1957). Failing to capture a prey in 
its initial attack, the cat does not continue the attempt. 

In the stomachs of cats caught while hunting, usually three to seven 
mice or voles have been found, but in "mouse" years significantly more. 
In Caucasian preserve on March 8, 1934, the stomach of one cat contained 
16 rodents and one tit [Parus]; their total weight was about 270 g. In the 
stomach of another cat 26 mice were found which weighed about 500 g 
(Satunin, 1915). 

Captive wild kittens aged 1.5 to 2.0 months are daily fed up to 10 mice 
each (about 180 g) while year-old animals receive an average of 900 g of 
horse meat every day (Teplov, 1938). 

Home range. Not much is known about the dimensions of the individual 
home range of a wild cat. In the Prut swamps a cat or a whole litter lived on 
every third or fourth floating island, comprising an area of 1.0 to 2.0 hectares 
(Lozan and Korchmar', 1965). In Caucasian preserve, in the proximity of 
the Kishinsk cordon, no less than 20 animals, or an average of one cat per 
60 hectares, were counted in the autumn-winter season of 1936/1937 in a 


total area of about 12 km^ (Teplov, 1938); in the Caucasus there are many 
regions where the population density of this predator is significantly higher. 

While living in a given section the cat leaves scent marks at different 
sites. The number of such marked sites greatly increiises during the period 
of estrus when the preanal glands of males and females enlarge and liberally 
secrete a compound containing trimethylamine and other odoriferous sub- 
stances. During the rut the male moves around trees with its tail raised and 
sprays its scent on the bark. Some cats begin to mark their territory when 
160 to 170 days old; the majority commence territorial claims only after 
365 days of age (Haltenorth, 1957). 

Burrows and shelters. In the Caфathians and Trans-Carpathians wild- 
cats live in the hollows of beech trees and in rocky fissures (F.I. Strautman; 
Konyukhovich, 1953). In the swamps of the Dnestr and Prut they live in 
summer in forks in the trunks of old willow trees, in abandoned heron 
nests, and in the hollows of willow trees and black poplars. In the Kochuf 
322 swamps (Moldavia) most cats live on floating islets. Usually the den with 
kittens is located in the central part of the island among densely grown reeds. 
Well-beaten pathways lead to it. Beside the den can be seen areas in which 
the kittens play and where prey is consumed (Brauner, 1914; Lozan and 
Korchmar', 1965). In the Caucasus, depending on the locality, these cats 
sometimes prefer to live in crevices among rocks, sometimes among hollows 
in trees, or sometimes in abandoned burrows of foxes and badgers (Dinnik, 
1914). In Caucasian preserve the most common shelters are hollows in the 
trunks of fallen trees, and more rarely hollows of growing trees, rock crevices 
and caves, and abandoned burrows of badgers and foxes. 

On May 11, 1936 in a preserve near the Kishi River the den of a cat 
was found on the bank of a small creek in a thicket of alder, nut trees, and 
wild apple. It was located in the hollow tmnk of a fallen rotten elm about 
45 cm in diameter. The hollow was about 30 cm in diameter and 7.0 m long. 
There was no bedding except the powdery wood rot. In the nest were the 
kittens (Teplov, 1938). 

In Zakatalo-Nukhinsk valley wildcats live almost exclusively in the 
abandoned burrows of badgers, which are numerous on slopes covered with 
beech forest (2 or 3 burrows per 10 hectares). In this same valley, at 
the upper forest boundary where burrows are few, cats use rock fissures for 
shelter. Only once was a den found in the hollow of a fallen poplar. It is 
characteristic that in places where the cat lives mainly in burrows, it does 
not climb trees to save itself from dogs but attempts to retreat into its burrow. 
On several occasions even when dogs chased a cat and began to attack it, 
the latter did not attempt to rescue itself by climbing a nearby tree (N.N. 

In taking up residence in the hollow of a living tree, the cat selects one 


which is not located very high above the ground. A hollow open above is 
not selected. On one occasion a wildcat's den was found in the center of 
a large firewood pile. The nest was made of dry grass and reeds and there 
were six very healthy kitteas with opened eyes (end of May, 1884; Rossikov, 
1887). It is evident that this cat does not dig a hole for itself. If the shelter 
is in a burrow or rocks, it is lined with dry grass, leaves, or bird feathers; 
while living in a tree hollow, the cat is satisfied with the wood dust present 
in it. It uses hollows or breaks as temporary shelters, or lies among dense 
vegetation. In years when the flea population is high in summer the cat does 
not live for long in the same den, but shifts from one to another. In winter, 
when soft and loose snow does not permit long-distant wandering, it remains 
for long times in the same den (Dinnik, 1914). In Caucasian preserve it 
uses the same site as a day rest for quite some time. In western Europe it 
selects hollows, rock crevices, and abandoned burrows of foxes and badgers 
for deas but sometimes makes an open nest in dense vegetation in marshes 
(Haltenorth, 1953). 

Daily activity and behavior. The wildcat mostly leads a crepuscular and 
nocturnal life. It usually sets out to hunt an hour or two before sunset and 
remains active throughout the evening and the first half of night, then rests, and 
reappears again only at dawn. In summer, however, it also hunts during the 
day. The stomachs of cats caught two hours before sunset contained freshly 
caught rodents. Many cases are known, when cats have stolen domestic fowl 
during the day and have been encountered by hunters. The wildcat is less 
cautious and feariess than other animals. It will permit a man on horseback to 
approach within 30 to 40 paces, after which it runs away unhurriedly. It swims 
well but avoids water when avoiding danger. A trapped cat rumbles and hisses 
at an approaching man but does not attack. A captive animal is very malicious 
and does not become accustomed to man; taming kittens is difficult indeed. 
According to observations made in the Caucasus, it invariably lives and hunts 
alone. According to data of eariier zoologists in western Europe, however, 
both parents live with the litter. Litters of young with two adult animals have 
reportedly been seen there. This information appears erroneous. The role 
of the male in raising kitteas is still not known (Haltenorth, 1957). 
323 Seasonal migrations and transgressions. These aspects are not known. 

In Caucasian preserve the wildcat does not, as a mle, wander even though 
food may be scarce (Teplov, 1938). In the Caфathiaas population changes 
are partly due to cats moving on during years of "poor harvest" of mice 
(F.I. Strautman). In western Europe cats descend from the mountains to the 
plaias in winters of abundant snow (Miiller-Using, 1963). 

Reproduction. In the Trans-Carpathians estms takes place from January 
to March (Konyukhovich, 1953); on the lower Prut (Moldavia) in early 
March (Lozan and Korchmar', 1965); in the northem Caucasus in Febniary 


(old calendar; Dinnik, 1914); and in Azerbaidzhan in the middle of February 
(Vereshchagin, 1942). However, according to recent observations based 
on captured pregnant females and the appearance of kittens, wildcats come 
into heat from December to February and May to July, and possibly in 
other seasons in different regions of the Caucasus. 

In Central. Europe animals in estrus are seen from the second half of 
February to the first half of March. Data are available which reveal a second 
breeding season, in autumn (Trauboth, 1961). In Scotland, between the end 
of May to early June, females often come into estrus a second time, and 
from September to October for a third time (Matthews, 1941). Females 
during repeated periods of estms have enlarged софога lutea from the first 
pregnancy, pointing to the possibility of two or three litters in a single year. 
Subsequent periods of estms are irregular. It is not known whether the mother 
suckles the kittens of the second, let alone the third litter, and whether this 
is even possible. Quite likely a subsequent litter appears due to the death 
of the previous litter. 

In some females estms continues for one to five days. Ovulation is 
induced [by copulation]. Spermatogenesis proceeds year-round (Haltenorth, 
1957). Mature sperm were found in the vas deferens of a cat caught on 
November 12, 1957, in the Ismaillinsk region of Azerbaidzhan Soviet Socia- 
list Republic (N.N. Rukovskii). 

During estms the preanal glands Ьесот^е enlarged in both sexes (see 
above) and sometimes vocalization, resembling that of domestic cats, is 
heard; both sexes are vocal. Fierce fights ensue between males during the 
mt. Gestation requires 63 days, sometimes up to 68 days (55 to 68 days 
in domestic cats). 

In Western Europe kittens of the first litter appear from the end of 
April to May. The second litter is seen at the end of July to early August, 
and the third (in Scotland) at the end of November to early December, and 
also mid-January (Matthews, 1941; Haltenorth, 1957). 

In the Trans-Carpathians kittens are born in April and May 
(Konyukhovich, 1953). In the Caфathians litters of four kittens with opened 
eyes were found in mid-May (F.I. Strautman; Tatarinov, 1956a); a juvenile 
cat with a body length of 30-40 cm was caught on November 15, 1952 
(Tatarinov, 1956). In Moldavia, on the lower Pmt, kittens are seen in 
the first half of May. On July 10 a cat was still suckling three grown-up 
kittens. On July 19 a young female had a body length of 63 cm, while 
a young male caught on July 25 weighed 1,900 g (Lozan and Korchmar', 

In the northern Caucasus kittens are seen toward the end of March 
or in April (old calendar; Dinnik, 1914) and in Azerbaidzhan in April (new 
calendar). Kittens with unopened and opened eyes were found at the end 


of May on the Malka River (Rossikov, 1887), and four kittens with eyes 
fully opened and milk teeth well developed were found on May 11, 1936, 
in Caucasian preserve. A female caught in Azerbaidzhan on February 23, 
1953, contained five large fur-covered fetuses (three females and two males). 
The fetuses averaged 117 mm in size and the weight of the females was 
68.6 g and that of the males 72.0 g. As these kittens would have been born 
toward the end of February, this cat must have mated in the second half 
of December. A female caught on February 21, 1953 in the same region 
had four implantation sites 43 mm in diameter and fetuses weighing 332 
mg. It evidently mated in mid-January and the kittens would have been born 
in raid-March. Lactating cats were caught on June 12, 1937 at Sevan 
(Armenia) and on June 21, 1939 in the Kubinsk region of Azerbaidzhan. 
A female caught on May 16, 1951 in the Zakatalo-Nukhiask valley contained 
324 five freshly implanted zygotes. Her milk glands were enlarged but devoid 
of milk (Rukovskii, 1955). 

Side by side with the spring appearance of kittens, some very late litters 
have been found in the Caucasus. On August 3 near Gelendzhik four kittens 
aged 10 to 12 days were found in the hollow of a tree (Obraztsov, 1928). 
In the 2!лка1а1о-КикЬ1п5к valley on August 15, 1950 a female contained 
seven fetuses with an average length of 68 mm each, indicating the last 
third of the gestation period. This cat would have given birth early in 
September. In the same region newborn kittens with a body length of 43 
to 49 cm and weighing up to 2,300 g were found in early August and 
September (Rukovskii, 1955). 

The number of kitteas in a litter in the Trans-Caфathians is three to 
six (Konyukhovich, 1953), in the Сафа1Ь1ап8 four (Lindemann, 1953; F.I. 
Strautman), in Moldavia three to five (Lozan and Korchmar', 1965), and 
in the Caucasus three to seven, most often four. Of the 12 litters studied, 
there were 4 kitteas in seven, 5 in two, and 7 in one. In hybrid litters of 
forest wildcat with domestic cat, there were no more than five kittens per 
litter (Haltenorth, 1957). 

In Western Europe the litter usually consists of three or four kitteas. 
The average for 41 litters was 3.3; one litter contained one kitten, 7 had 
two, 15 had three, 14 had four, and 4 had five each. In Scotland a litter 
contains two to four kittens (Haltenorth, 1957) and in France two to six 
(Beaufort, 1961). 

Growth, development, and molt: Kittens are bom blind and helpless. 
They gain sight on the 9th to the 12th day. At that time they are covered 
with a fuzzy co'at; the top of the body is very pale yellow, with dark brown 
spots fusing on the back into broad dark bands. The hind legs are mottled 
with numerous narrow stripes. Five broad dark rings occur on the tail and 
the tip is dark. The sides of the head, in the region of the ears and cheeks. 


have a reddish tinge. A reddish spot is present on the nose. Some dark stripes 
traverse the top of the head and the tips of the ears are dark. The underside 
of the body is very pale yellow. The neck and underside of the chest are 
sprinkled with faint dark-colored spots. These kittens differ from domestic 
ones in their larger, more broadly opened ears. 

The weight of four newborn kittens in the Cracow Zoological Garden 
was 38, 42, 44, and 45 g. Growth and development of kittens of European 
wildcats proceed rapidly (Lindemann and Rik, 1953; from Haltenorth, 1957): 

Age (days) Weight (g) Age (days) Weight (g) 
























Incisors erupt at the age of 14-30 days and milk teeth are replaced 
by permanent ones at the age of 160-240 days. Sexual maturity is attained 
in 300 days. The young begin to hunt with their mother at the age of 60 
days and travel with her up to 140 to 150 days, after which they move 
independently. The first encounters with solitary young cats which have 
left their nest occur in mid-June in Caucasian preserve (Teplov, 1938). At 
this time they leave their mothers and hunt for food on their own. By August 
young cats hunting singly are a common phenomenon. 

Lactation continues for three to four months but kittens begin to eat 
meat also (mice, young hares, young rabbits, and small birds) at the age 
of 1.5 months. When slightly over a month old they set off from the den 
and play, often climbing trees. Trees soon become the favorite places for 
play and serve as refuges when danger is near. 

By the end of September in the Caucasus the young of spring litters 
are only slightly smaller than adult females in appearance and weigh over 
2.0 kg. In Caucasian preserve a male caught on September 10, 1936, weighed 
2,235 g and a female caught on November 19, 1936, weighed 2,800 g; a 
female on December 25, 1936, weighed 2,500 g and a female on January 
22, 1937, weighed 1,900 g. 
325 As long as her kittens are small the mother is extremely restless, does 
not leave them alone for long, protects them from attack by small predators, 
and in the event of danger, transports them to a new den. 

The maximum longevity of the wildcat is 21 years; usually it survives 
for 13 to 14 years (Shortridge, 1934). The report that one cat lived for 39 
years (Pidoplichko, 1956) is not reliable. 


Cats of the silvestris group molt twice a year. Spring molt ends in May 
and autumn molt in mid-November. The [winter] coat is fully developed 
from mid-November through March. 

Enemies, diseases, parasites, mortality, and competitors. Enemies of 
wildcats are almost unknown. In most samples of food and feces of the 
wolf in the Caucasus, remains of wildcats have not been found. Only one 
predatory battle has been witnessed. After a stubborn stmggle a forest marten 
overcame a wildcat, tore open its abdomen, and ate its liver (Satunin, 1915). 
However, considering that the cat usually lives at places where there are 
rocks and tall trees for refuge or dense thickets and abandoned burrows, 
serious enemies can hardly exist. In Central Europe many young cats are 
killed by marten (Trauboth, 1961). 

The diseases suffered by wildcats have not been studied. In zoological 
gardens they sicken and die from infectious enteritis (Tsvetaeva, 1949). 

Wildcats are highly parasitized by helminths. Of ten cats examined in 
Caucasian preserve, eight were infected with up to 70 nematodes, and between 
6 and 73 cestodes were found in all ten (Teplov, 1933). In some cats in Georgia 
five species of helminths have been identified: Hydatigera taeniaeformis, 
Diphyllobothrium mansoni, Toxocara mystax, Capillaria feliscati, and 
Ancylostoma caninum (Rodenaya, 1951). In Azerbaidzhan cats are infected 
by Hydatigera krepkogorski and T. mystax (Sadykhov, 1955). In the Trans- 
CaucasiLs the majority of cats caught were infested with the tick Ixodes ricimis. 
In some summers cats there are severely infested with fleas of two species: 
Ceratophyllus consimilis and Ceratophyllus sp. The first has been found in 
large numbers in Norway rats living in the forests (N.N. Rukovskii). 

The competitors of wildcats are jungle cat, jackal, fox, marten, and 
other predators. The jungle cat is a particularly serious competitor. However, 
the two cats usually occupy different biotopes. In the Caucasus jungle cats 
inhabit the lowland sections while forest wildcats reside higher up along 
the mountain slopes in beech forests. Usually at places where wildcats are 
numerous jungle cats are few or altogether absent, and vice versa. In the 
Lenkoran lowland jungle cats are numerous and wildcats absent. Sometimes 
such a distribution can be explained by foilurc of the ranges to meet in 
a small territory. 

Population dynamics. This aspect is almost unstudied. In the Car- 
pathians, following a year of a rich harvest of beechnuts, mouse-like rodents 
proliferate and in roughly one year's time the cat population likewise shows 
a significant rise. The reverse correlation has also been noted — a decrease in 
cat population in years when the rodent population is low. As a result of 
fluctuations in the cat population the number of skins tanned annually varies 
more than twofold (F.I. Strautman). The cat population increases in alternate 
years or at intervals of two or three years. Evidently in the Caucasus, 


326 Fig. 192. Footprints of a wildcat on sand. Match box for relative scale. 

Azerbaidzhan. December, 1957. Photograph by N.N. Rukovskii. 

population shifts in wildcat depend on the availability of "mice," which 
are particularly abundant after a good crop of beechnuts and acorns. The 
cat population should decrease significantly in unusually snowy and 
protracted winters. In such winters it becomes difficuh for these predators 
to obtain food and they suffer from starvation (for example, in the winter 
of 1953/1954 in the Kuban). Cats living in the swamp forests of the Dnestr 
quite often die not only in spring but also in summer floods (Brauner, 1914). 
Floods are fatal to kittens in the swamps of the Pmt, Danube, Kuban, Terek, 
and valleys of other rivers. Finally, the population may fluctuate as a result 
of epizootic diseases. In recent years in Azerbaidzhan the cat population 
has decreased considerably due to intensive shooting (N.N. Rukovskii). 
326 Field characteristics. Generally bigger than domestic cats. Their color 

is monochromatic, mottled with dark transverse bands which are sometimes 
barely distmct. The tail is furrier than that of domestic cats and looks 
chopped; there is a light-colored spot on the throat. Wildcat differs from 
jungle cat in variegated coloration, and a much longer, and, chiefly, in a 
more luxuriant tail. The tracks resemble those of domestic cats but are bigger 
(Fig. 192). Generally near trails leading from a wildcat's lair broken reeds 
can be found, on which the cat has cleaned its claws; on the trails themselves 
lie relatively large-sized fecal pellets with a blunt end. This cat eats only 


the internal organs of large prey: lungs, heart, and liver. Its voice is similar 
to that of domestic cat but coarser. (A.S.) 

Practical Significance 

A fur-bearing animal, but the skin is of little value; the average price after 
tanning is about 50 kopecks. Pelts play only a small role in skin-processing 
plants in the Trans-Caucasian and Caucasian republics. 

The output of tanned skins in recent years can be judged in part from 
the data given in Table 15. One should keep in mind, however, that tanned 
skins of wild and other cats are classed under the general name "wildcat" 
and hence only information about the catch of "wildcats" in the Ukraine 
and Moldavia may be attributed entirely to the forest wildcat. 

The All-Union standard for pelts of wildcats refers to the group 
"wildcat," which is further subdivided into "forest". 

When living close to cultivated areas wildcat is useful in destroying 
rodents (see above). In most regions, however, it is a pest of game, killing 
nutria, muskrat, hares, pheasants, partridges, and waterfowl.'*-'* In the 
327 Caucasus this cat, along with jungle cat and jackal, is considered a major 
hindrance in nutria husbandry. It has long been a serious pest of muskrat 
breeding on the Prut, Kuban, and Terek. At some places it also causes 
damage to poultry farming. 

Table 15. Tanned skins of "wildcats" (total) in the Soviet Union 
system of Zagotzhivsyr' [raw materials procurement] 

Region of procurement 





Ukrainian SSR 





Moldavia SSR 





Azerbaidzhan SSR 





Georgia SSR 





Armenia SSR 





Northern Ossetia 





Stavropol ' 





Groznensk district 





Dagestan ASSR 





Kabardino-Balkarsk ASSR 





Krasnodarsk territoi7 





Total for the Soviet Union 





''^The "sanitary" role of this carnivore should also be taken into consideration. By des- 
troying mostly weak and diseased animals, it "improves" the fitness of the game population. 


Special hunts for wildcats were not practiced in the past. Many were 
caught in traps and other snares set for marten, and by shooting, using dogs 
to tree them. Following their intense persecution in Trans-Caucasus as a 
pest of game, they are now caught in traps placed on their pathways without 
a bait, or at abandoned burrows of badgers and foxes. The pelts are used 
in ladies' fur coats and sometimes converted into imitation sealskin by 
cropping and dyeing them black, but the color of such dyed skins is more 
variable than in other furs. 

While the forest wildcat is one of the many predators in the Caucasus 
and Trans-Caucasus, it is now extremely rare in the western parts of the 
range within the Soviet Union (except some western regions of the Ukraine). 
In Belorussia, the Ukraine (except the Carpathians), and Moldavia, it is 
an interesting monument of nature and should be conserved there by banning 
hunting until its population is restored. Hunting wildcats is already banned 
in the Trans-Carpathians and other western regions of the Ukraine. The 
small population of this cat in Soviet forests poses very little danger to 
game; on the contrary the cat has become a favorite object of hunting. (A.S.) 


1780. Fells lybica^^ Forster. In Bujfon. Naturg. vierfuss. Thiere, vol. 6, 

p. 313. Gafsa, Tunisia. 
1791. Felis ocreata. GrciQlm. Anhang zu Bruce. J., Reisen in Africa, vol. 

2, p. 27. Abyssinia. 
328 1830. Felis ornata. Gray. Illustr. Indian ZooL, vol. 1, pi. 2. India. 
1834. Felis servalina. Jardine. Nat. Library, Felinae, p. 232. India. 
1874. Felis caudata. Gray. Proc. Zool. Soc, London, p. 31, pi. 6. 

Dzhanydar'ya by southwestern part of Aral Sea (northern border of 

Kyzylkum). Not Kokand and not Bukhara! (Birulya, 1912). 
1876. Felis shawiana.^^ Blanford. Journ. As. Soc. Bengal, vol. 45, p. 49. 

Yarkand, Chinese Turkestan. 

"'''Also called spotted, and long-tailed cat. 

"•^Evidently a typographical error by the author. Usually the correct classical spelling 
(libyca) is used. 

''^Pocock (1939) stated that in the material used by Blanford for his description, the pelts 
belonged to steppe cat and the type was designated as a skull of F. chaiis; hence the name 
shawiana should be placed among the synonyms of jungle cat. This view has been accepted 
by some zoologists (Ellerman and Morrison-Scott, 1951 and 1966). Usually (Ognev, 1935; 
Haltenorth, 1953; Weigel, 1%1) the name shawiana is used for steppe cats of eastern Turkestan. 
This is the correct usage, further supported by the fact that (previously ignored) jungle cats 
in the east do not occur farther than Syr-Darya (see "Geographic Distribution" of this species) 
and have not been known for eastern Turkestan. Steppe cats are widespread there (Murzaev, 
1966). The name shawiana is the first among the series of names assigned to cats of Central Asia. 


1914. Felis caudata morpha shnitnikovi. Birula. [=] Birulya. Ann. Mus. 

Zool. Ac. Sc. Petrograd, vol. 19, p. 11. Semirech'e. 
1914 (1915). Felis caudata griseoflawa. Zukowsky. Arch. Naturg., A 80, 

9, 95. Upper course of Chu. 
1914 (1915). Felis caudata longipilis. Zukowsky. Ibid., A 80, 9, 97, District 

south of Balkhash. Nomen praeoccupatum — F. longipilis Fitzinger, 

1868 (tiger). 
1914 (1915). Felis caudata macrothrix. Zukowsky. Ibid., A 80, 10, 127. 

New name for F. с longipilis Zuk., 1914. 
1914 (1915). Felis (Felis) murgabensis. Zukowsky. Ibid., A 80, 10, 127. 

Takhta Bazar on upper Murgab. 
1914 (1915). Felis caudata matschiei. Zukowsky. Ibid., A 80, 10, 130. 

Foothills of Kopet-Dag at Geok-tepe west of Ashkhabad, 

1914 (1915). Felis (Felis) maimanah.^^ Zukowsky. Ibid., A 80, 10, 130. 

Maimene, northern Afghanistan. 
1930. Felis ornata issikulensis.^^ Ognev. Zeitschr. F. Saugetierk, vol. 5, 

p. 67. Northwest shore of Issyk-Kul [Lake]. 
1945. Felis ornata araxensis. Sarkisov. Dokl. AN Arm. SSR, 3, 3, 88. 

"Lowlands of southeastern Trans-Caucasus"; more precisely (p. 89), 

steppe at Arazdayan on the Araks (about 45° E. Long.). (V.H.) 


In general appearance steppe cat is very similar to forest wildcat, but appears 
somewhat smaller and lighter. Tail relatively slightly longer (usually more 
than one- half length of body) and, more importantly, thin, and not truncated, 
but pointed at the tip. Comparative lighter color of steppe cat due in part 

"•^Reference to the type locality has been corrected. Usually it is shown as 110 versts 
south of Geok-tepe in the Trans-Caspian district at 38° N. Lat. and 57° 30' E. Long, in the 
Kopet-Dag. The point 110 versts south of Geok-tepe falls far inside Iran, deep in the mountains, 
roughly midway between Geok-tepe and Sabzevar, and very far from the foothills of the 
Kopet-Dag lying in the immediate proximity of Geok-tepe. Latitude 38° passes right by 
Geok-tepe atong the northern foothills of Kopet-Dag. 

^^Zukowsky bestowed this name based on a description of skins oi Felis caudata from 
Maimene published by Scalli (1887). Ellerman and Morrison-Scott (1951 and 1966) rather 
uncertainly (with a question mark) assigned it to F. chatis. However, according to other 
researchers (Ognev, 1935; Haltenorth, 1953), it is quite cleariy a description of skins of steppe 
cat. A skin preserved in the Calcutta Museum (India), excellently described by Dr. B. Biswas 
(Zoological Survey of India), has not been preserved. This zoologist also thought that the 
skin from Maimene belonged to a steppe cat (F. libyca). (V.H.) 

^^The form has been described from a feral domestic cat and from hybrids of wild 
F. s. caudata and domestic cats. Material in Zoological Museum, Moscow University. 


330 to the fact that, even in winter, its coat does not attain the levels of density, 
length, and luxuriance seen in forest wildcat. Hair on tail much shorter in 
the former and not so dense, luxuriant; close-fitting, hence tail appears much 
thinner. -'^^ Bare section on nose not flesh-colored (pink) but black or dark 
(Fig. 193). 

"Pelage not long, fairly soft, and not sparse. On the back about 3,500 
hairs occur per cm^, with eight of underfur to every top hair. Hair on 
abdomen longer than on the back (abdominal type of coat) and guard hair 
stands out prominently from top hair; various categories of top hair (four) 
do not differ greatly in length, and underfur equal in length to hair of category 
IV. Differences in hair thickness are considerable, however. Length of guard 
hair on back 51 mm and thickness 92 microns; corresponding values on 
abdomen 58 mm and 59 microns. Length of top hair on back 43, 39, 37, 
and 35 mm and thickness 102, 76, 56, and 42 microns; corresponding values 
on abdomen 54, 49, 46, and 44 mm, and 48, 42, 35, and 31 microns. Length 
and thickness of underfur on back 35 mm and 23 microns and on abdomen 
38 mm and 21 microns" (B.F. Tserevitinov). From the foregoing description 
it is obvious that the hair coat of steppe cat differs greatly from that of 
forest cat (Caucasian) in type of fur. 

The main features of coloration of the steppe cat are quite specific but 
its details diverse and variable. A detailed description would be quite com- 
plicated therefore, and only the main features are described. -'^^ 

General background color of the skin in the upper part of the body 
very light (Fig. 194). In darker varieties, a fairly pure light gray color; pale 
yellow or sandy shade absent. In many varieties skin steadily fades and 
acquires a yellowish or even sometimes distinct yellow tone. When the 

331 color is highly faded, the background is a very pale white with a light 
creamy-yellow (pale yellow) tone. Along the middle of the back (along 
the spine) the main color is usually somewhat darker, forming a more dark 
gray, brownish, or ocherous band corresponding to the general background 
but varying in different forms. In some animals, mainly the extremely light 
colored ones, aarkening on spine very faint or almost not visible. 

Entire upper body covered with small, round, solid spots. Their diameter 
in different parts of the body differs, but only insignificantly and is associated 
mainly with length and density of pelage in a given region. At places (back) 
where the coat is very short and dense the diameter of the spots is less than 

^^It is possible that some differences in body proportions do occur between forest and 
steppe cats, tlie latter having somewhat larger heads. 

^^Description based on material from Turkestan and southern Kazaklistan (about 70 
winter skins in the collection of MMU). It differs significantly from the description commonly 
found in literature, given by S.I. Ognev (1936). This author had only a small sample of material 
and exaggerated the geographic importance of some features. 




330 Fig. 194. Adult male steppe cat in winter coat. Aksu-Dzhebagly preserve. Talassk 

Alatau, western Tien Shan. January, 1953. Photograph by F.D. Shaposhnikov. 

on the abdomen where the coat is longer and more sparse. This feature 
also affects the shaфness of the outline of a spot, but only minor individual 
variations are seen. The nature of the spottedness evidently also depends 
on age — small-sized skins apparently appertain to newborn kittens, which 
have the smallest and also densest spots. 

On the whole the spotted pattern is highly variable, due to various 
combinations of features (Fig. 195). The spots may be relatively large or 
small, shaфly contoured or with weakly or greatly blurred outline, and 
disposed densely or relatively sparsely. The color of the spots in most cases 
is black, sometimes pure bfaek, and sometimes slightly brownish. In animals 
with a sandy-yellow background the spots are a stronger brownish shade. 
The spots are neither arranged in clusters nor as rosettes. They are circular 
but in some animals along the middle of the back (along the spine), especially 
the rear portion, but sometimes also between the shoulders, they are some- 
what elongate. The size of an individual spot generally remains constant. 
In extreme case (rare) the spots may form two short longitudinal parallel 
rows on a short segment in the sacral region. 

Usually the spots exhibit no tendency to form transverse rows or 




transverse stripes on the trunk. The "tiger pattern," consisting of solid 
transverse stripes, is also never found.-'''^ "Transverse rows" of spots, 
reported by different workers (Ognev, 1935; and others), are quite indistinct 
and strictly speaking imaginary; only in a few skins are they distinct. Two, 
sometimes three, distinct solid, dark (black), transverse stripes are generally 
present on each side of the shoulders as well as behind them. Along the 
spine spots do not coalesce. Distinct solid stripes are seen on the thighs, 
which are longitudinal or oblique-longitudinal in disposition. Sometimes 
there are no stripes on the thighs. 

Color is also variable between the shoulders and along the top of the 
neck. In one extreme case a very dark uniform field with an indistinct pattern 
was found here; in another skin there were fairly distinct (some-times very 
definite), narrow, black or blackish-brown, longitudinal stripes. They may 
be quite sharp only in some sections, or throughout, and usually number 

The color of the underside is light, mainly white with a light gray, 
creamy, or pale yellow tinge. Its intensity and hue depend on the general 
color of the upperside. Spots, far more blurred and often much larger than 
on the back, occur only on the chest and anterior part of the abdomen; the 
posterior part of the abdomen, inguinal region, and inner surface of the 
thighs are devoid of spots and very light, almost white. The region between 
the forelegs, lower neck and throat up to the chin, and the lower J ins are 
without spots, or occasionally with a few indistinct spots. These '■'"gions 
in some animals are pure white and in others a very famt shade of ^Ьс color 
corresponding to the basic tone. In many animals, from both sidf^^ of the 
base of neck, a very light-colored patch descends in the form ot a cape; 
it usually forms a broad band with indistinct contours. In some, on the 
333 neck and throat above this band or as an anterior continuation of the band, 
an intense ocherous shade is present, and there are occasionally fairly bright, 
often irregular spots of the same color on the white field of the throat. 
Most of the tail is the same color as the general color on the back. 
Along the upper two-thirds (or thereabouts) of its length a black or dark 
brown, narrow stripe occurs. The very tip of the tail is black; in the region 
in front of the tip there are two or three, more rarely four or even five, 
black transverse rings. Thus the colored portion may occupy slightly more 
than one-third the length of the tail. The upper parts of the feet are mono- 
chromatic, pale yellow, and the lower parts dark brown, sometimes black. 

The head exhibits a fairly complex pattern (Fig. 196). The upper lips 

^^Distinct transverse stripes (apart from those on the shoulders) and more so the "tiger 
pattern" observed in cats caught in the wild constitute a feature of hybrids obtained from 
crossing with domestic cats. 


Fig. 196. Adult male steppe cat. Hi delta. Semirech'e. February, 1967. Photograph 
by Yu.V. Vladimirov. 

are light, pale yellow-white and a ring of similar color seen around the 
eyes; ocherous sections occur along the sides of the bare portion of the 
nose and on its upper side where there are short hairs. The region between 
the eyes, the space between the eye and ear, and the field under the ear 
are monochromatic, gray, and fairly intense. The top of the head is covered 
with a dark gray coat but the tips of the hairs are light-colored. Dark- and 
light-colored longitudinal bands absent on the head or very faintly 
discernible. Sometimes on the forehead and top of head dense clusters of 
brown spots occur but usually there is no pattern whatsoever. Narrow, most 
often pure black stripes, usually four in number, occur, which, as mentioned 
earlier, sometimes run along the neck into the region between the shoulders. 
From the comer of the eye under the ear, along the rear of the cheek, a 
dark black or brownish narrow stripe extends to the level of the rear edge 
of the base of the ear. Another similar band, commencing under the eye, 
extends back below the former to the same level or slightly beyond. In 
general, the length of both stripes is variable but they are always present. 
Sometimes thin, short, additional bands occur under the eye in front. The 
outer side of the ears is light gray, darker along the anterior inner rim, and 
lighter at the base, close to the outer rim. The inner surface of the ear is 


covered with long white hair. The tips of the ears terminate in thin tufts 
of black hair 15 to 18 mm long. The color of the head, as also the pattern 
of markings, is highly variable (Fig. 197). 

334 Sexual color dimorphism absent. Summer coat distinguished by a 
slightly lighter background color and more sharply contoured spots. Young 
animals, until they attain their first winter coat, exhibit very small but dense 
spots on a fairly soft fuzzy coat (Fig. 197). The general shade of the coat 
is very light. Much smaller but dense spots, though not as dense as in kittens, 
are evidently preserved even in the first winter coat (see above). 

In the category of exotic variations, chromists and partial albinos are 
known. The latter might be hybrids with the domestic cat. 

The skull in its various features is similar to that of the skull of the 
silvestris group, but differs significantly in the structure of the auditory 
bulla'^-'' (Fig. 29). It is somewhat larger, quite voluminous, and bulges 
uniformly. It mainly differs in the larger volume of the ectotympanic 
chamber. As a result the section between the ear and the Eustachian openings 
is more roomy and the anterior boundary projects noticeably forward. It 
is shifted much more forward from the opening of the Eustachian tube than 
the boundary of the section lying inside. In the skull of forms in the silvestris 
group, both these boundaries fall at the same level in relation to the sagittal 
plane, or the boundary of the inner plane is shifted forward even slightly 
more. The anterior margin of the ectotympanic chamber in the steppe cat 
lies at the level of the anterior margin of the postglenoid process or almost 
at this level; in the forest wildcat it is significantly shifted posterioriy and 
situated behind the posterior margin of the postglenoid process. 

The opening of the external auditory meatus in the steppe cat is large 
in absolute and relative terms, its size increasing in relation to the increase 

335 of the ectotympanic chamber. The relatively large size of the external 
auditory meatus is not obvious because of the overall size of the auditory 
bulla (profile); furthermore, in the steppe cat the maximum diameter of 
this opening is greater than the length of the crown of the third upper 
premolar; in the forest wildcat it is smaller or equal to it. Such a structure 
of the auditory bulla of the steppe cat represents a distinct adaptation to 
desert conditions, totally homologous to such adaptation in many other 
mammals (carnivores and rodents). 

There are no differences in dentition but the peak on the anterior- 

■^As pointed out above, some workers (Pocock, 1951) assume that there are no dif- 
ferences between the skulls of F. silvestris and "F. libyca," and that these "species" differ 
only in color. Other characteristics have been verified on a large number of animals in both 
groups, but only from within the Soviet Union. Evidently these characteristics are common 
to animals of adjoining territories. Material from Africa has not, however, been investigated. 
The form sarda ("transitional") has auditory bulla of the forest wildcat type. 


334 Fig. 197. Young male steppe cat. Badkhyz presei^ve. Southern Turkmenia. May, 

1962. Photograph by Yu.K. Gorelov. 

inner ciisp of the upper camassial tooth is evidently somewhat better devel- 
oped and sharper. Both upper premolars were present in 667c of the animals 
examined (in 30 of 45 males and females taken together), both were absent 
in 18% (8 of 45), or one absent in 15 to 16% (7 of 45). In males (22) both 
teeth were present in roughly 64% (14), both absent in 27%: (6), and one 
absent in 9.0% (2). In females (12) both premolars were present in 66%- 
(8), both absent in 8.0% (1), and one absent in 25% (3) (material from 
Turkestan). These ratios are somewhat different than the corresponding ones 
in the Caucasian forest wildcat. 

Sex-related and age-related skull changes correspond to similar 
changes in the forest wildcat. Geographic variation has been noticed but 
not thoroughly studied. It is not seen within the boundaries of the Soviet 

In body and skull dimensions forms representing the group correspond, 
on the whole, to forest wildcats of Europe and the Caucasus but on the 
average are smaller; some southern animals are significantly smaller. In any 
case the maximum size and weight attained by forest wildcats of Europe 


are not attained even by members of the biggest subspecies of steppe 
cats.-''^ This pertains also to the Turkestan form (see ''Geographic 
Variation"). Females are perceptibly smaller and lighter than males. 

Within the species as a whole Bergmann's rule is distinctly manifested 
(biggest cats found in Europe and Middle Asia); the same is true of Gloger's 
rule — most richly colored cats found in Europe and the Caucasus among 
them especially in Scotland — the whole of the desert-steppe members of 
the libyca group are pale; some desert forms of Africa are particularly pale 
and exhibit a much reduced spotted pattern. Allen's rule is manifested in 
body proportions. These rules often apply within each group of subspecies 
individually. (V.H.) 

Origin of Domestic Cats 

At present it is thought (Pocock, 1951; Haltenorth, 1953 and 1957; Zeuner, 
1957) that the ancestral form of domestic cat was the North African bay 
or Libyan cat, F. (F.) s. libyca Forst. It was domesticated in Egypt very 
long ago. According to some authors the first reference to domestic cat 
dates back to about 4000 B.C., or to the First Dynasty, about 3000 B.C. 
By 2000 B.C. at the beginning of the Middle Kingdom, cats were already 
quite common in Egypt as domestic animals and widely distributed. 
According to other authors (Zeuner, 1967) portrayals of cats in the Middle 
Kingdom represent wild animals and it wiis only in the New Kingdom, 
about 1600 B.C., that the cat became a domestic animal, i.e., domestication 
occurred not very long ago.-""^ 
336 Initially domestication of cats arose from the need to combat house 

and granary rodents. Mystic or religious importance was assigned to the 
cat later. It was even considered a sacred animal (cult of goddess Bast at 
Bubastes). Bodies of cats were mummified and preserved in large numbers. 
A study of a large number of these mummies (Morrison-Scott, 1952) revealed 

■'^^his refers to norma! variation. As pointed out above, especially large wildcats of 
western Europe (weighing from 11-15 to 18 kg), if these figures are not erroneous or 
exaggerations, probably represent hybrids with domestic cat, i.e., an exotypical phenomenon 
Hke the "giant" Amur cats. Hybrids of steppe and domestic cats do not attain such "giant" 
sizes; at least they are not known to occur within the Soviet Union. 

^^B. Brentjes (1965) assumes that the cat was domesticated in the Near East much 
earlier than in Egypt. According to him, representations of cats in Erihon and the Jordan valley, 
from the end of 6000 and early 5000 B.C., depict domestic animals, as do representations 
from southern Iraq from early 300 B.C. Statuettes of cats from Kachilar in Asia Minor date 
to 6000 B.C. Thus the Libyan cat, F. s. libyca, is not the only ancestor of domestic cat; 
it had already hybridized with the Middle East cat. This view has found no confirmation 
(Zeuner, 1967). Brentjes has not taken into consideration that forest wildcats lived in the 
Kachilar region, which removes the entire question of ancestry to a different plane. (V.H.) 


that F. s. libyca was domesticated and, furthermore, long before our time 
the Egyptians had a special domestic form of cat, which was slightly bigger 
than the wild one ("F. bubastis'" Hempr. and Ehrenb., 1832). 

The possibility should not be completely excluded of other pockets 
of domestication in antiquity, independent of Egypt, in the agrarian states 
of the Near and Middle East, and India, i.e., in the range of the subspecies 
of group omata-caudata, primarily in Persia and Mesopotamia. However, 
such a probability appears dubious and evidence quite inadequate. In these 
countries the cat more likely arrived from not-so-distant countries such as 
Egypt. Ancient China could not have produced the domestic cat because 
the species silvestris -libyca was absent there. The domestic cat of China 
came from the Middle East through India just before our era (not earlier 
than the sixth century). Europe was also not the home of cat domestication 
and the form F. s. silvestris cannot be included among the ancestors of 
domestic cats. Not only the low population density and lower cultural level 
of the inhabitants of Europe, but also the wild nature of the forest cat, exclude 
this possibility. The forest cat does not tame as readily as the steppe cat 
and is far less suited to domestication. The cat in Europe either came from 
Egypt directly or via the Near East. 

Evidently there were two routes of range expansion — via Italy (Rome) 
or via the Balkans and Black Sea region. The cat appeared first in Europe 
in the Black Sea region (Ol'viya, sixth to the first century B.C.; PidopUchko, 
1952). Reports of its appearance in Rome are dated much later; the first 
reference pertains to the fourth century A.D. The cat colonized central and 
western Europe only in the Middle Ages and was a rarity in the far west 
(England) even in the tenth century. 

Colonization of cats in Russian territory occurred roughly in the same 
epoch as in central Europe. As indicated by the period in which some finds 
are reported and the overall political and cultural situation obtaining in 
ancient Russia (relations with Byzantium in particular), the cat evidently 
came into Russia not only from the west but also from the south. Its remaias 
are known from the seventh and eighth centuries A.D. in the Sum district 
(Volyntsovo Putivl'sk region), from the eleventh to the thirteenth centuries 
in the Zhitomir district, and from the twelfth and thirteenth centuries in 
Kiev and elsewhere. To the north remains of domestic cats have been found 
in horizons of the seventh to the ninth centuries in Staraya Ladoga, tenth 
to the thirteenth centuries in Moscow, eleventh century in Novgorod, and 
thirteenth and fourteenth centuries in Pskov and elsewhere (Pidoplichko, 
1956; Tsalkin, 1956). 

As mentioned earlier, while discussing hybridization of domestic and 
forest wildcats, a change in landscape and a decrease in population of forest 
cats could have led to instances of hybridization with domestic cat. However, 


the influence of domestic cat on wild was greatly exaggerated until just 
recently. It was thought, and sometimes still is, that domestic cats exerted 
a major influence on wildcats. The picture in fact was otherwise. Wild forest 
cats evidently exerted some influence on European domestic cats. Animals 
with a tiger-like coat (transverse stripes or distinct vertical rows of spots 
on the body) exhibit a pattern characteristic of forest cats of Europe. Even 
in Turkestan this pattern is found to varying degrees only in hybrids of 
steppe and domestic cat and not seen among "pure" wildcats (F. s. caudata). 
In general, however, the influence of European forest cat on domestic cat 
was very insignificant. 

It has been suggested that the Angora cat (also called Persian cat in 
the west) evolved as a result of hybridization of domestic cat with the 
Caucasian forest cat in Asia Minor. Its long coat was apparently acquired 
from wildcat together with some other features of its "face" and general 
appearance (Pocock, 1907; Schwangart, 1932; and other earlier authors; 
337 Haltenorth, 1953). This suggestion has not been confirmed. The lone 
argument is based on the assumption that this form originated in Asia Minor 
(Angora-Ankara) where the forest wildcat is present. Such a view is hardly 
justified; long-haired breeds of domestic cats are also known in Middle 
Asia ("Bukhara"), Siberia ("Siberian"), and China. Evidently these species 
are also products of selection like other long-haired "Angora" animals 
(rabbit, goat, and guinea pig) and Pekinese dogs. 

In discussions on the origin of domestic cats certain other views have 
been expressed which are no longer accepted. Thus, even Pallas (1811) 
suggested (some authors have interpreted his suggestion as an affirmation) 
that the Angora cat is derived from the manul cat since "domesticated manul 
cats readily mate" with domestic cats ("licet mansueta cum hoc facile 
coeat"). Pallas' suggestion had to await confirmation for want of anatomica